Signal-centered action patterns of dogs in appetitive classical conditioning

The movements of five unrestrained dogs were monitored during discriminative Pavlovian conditioning. An auditory-visual stimulus from one source (CS+) was followed by meat morsels (US); the same stimulus from another source (CS?) was not. Sources were equidistant from the site of US delivery. Before each trial, animals were required to position themselves at a starting location equidistant from both sources and removed from the US site. The stable behavior pattern in most subjects included approach to and contact or near contact with CS+, followed by approach to the US site. Dogs showed individually distinctive action patterns to CS+, in some cases suggestive of soliciting, in another of sight-pointing. Similar actions were not evoked by CS?. In postacquisition tests, patterns were generally unaffected by increased deprivation or by relocating the starting position next to the US site. The auditory component of CS+ was more effective than the visual. With a response-contingent procedure animals were successfully trained to approach the US site directly, although many trials, and in one case a special procedure, were required. It was proposed that the experimental signaling arrangement mimics a natural signaling sequence and induces appetitive behavior to the CS that corresponds to the behavior induced by certain natural signals of the US.     

Cerebellar norepinephrine modulates learning of delay classical eyeblink conditioning: evidence for post-synaptic signaling via PKA

The neurotransmitter norepinephrine (NE) has been shown to modulate cerebellar-dependent learning and memory. Lesions of the nucleus locus coeruleus or systemic blockade of noradrenergic receptors has been shown to delay the acquisition of several cerebellar-dependent learning tasks. To date, no studies have shown a direct involvement of cerebellar noradrenergic activity nor localized the post-synaptic response to cerebellar beta-noradrenergic receptor signaling. Using ipsilateral, localized infusions into cerebellar lobule HVI and interpositus (IP), we have established that blocking beta-noradrenergic receptors with propranolol significantly impairs acquisition of conditioned responses. Furthermore, interrupting activation of cAMP-dependent PKA in the cerebellum using Rp-cAMPS completely prevents acquisition. However, neither blocking beta-adrenergic receptors nor blocking PKA activation significantly interferes with performance of established conditioned responses when administered after the learned response is formed.     

Effect of interpolated temporal activity on temporal short-term memory

The purpose of this research was to determine the retroactive interference effects of a single interpolated task (i.e., one temporal duration) on the retention of a criterion duration. This research is of interest because the mnemonic structure of temporal information of different durations is uncertain. Previous research has indicated that there might be a difference in structure for durations of 1 and 4 sec., although the results are inconsistent. Thus, two criterion durations of 1 and 4 sec. and five interpolated durations (i.e., 60%, 80%, 100%, 120%, and 140% of the duration of the criterion) were utilized under the method of reproduction. In addition, subjects were instructed to use either a counting strategy or none (referred to as conscious time estimation) to facilitate the retention of the temporal information. Recall was less variable when using a counting strategy than not and when estimating 1 sec. than 4 sec. However, there was no effect of interpolated activity when comparing performance across different interpolated conditions (no interpolated activity). Apparently, one interpolated duration is not sufficient to produce structural interference with a single criterion duration.     

Inhibition of monoADP-ribosylation prevents long-term memory consolidation of a single-trial passive avoidance task in the day-old chick

The cytosolic posttranslational protein-modifying mechanism of monoADP-ribosylation has been implicated in long-term potentiation, a synaptic model of memory formation. The current study investigated the effect of inhibiting mono(ADP-ribosyl) transferase on memory for the passive avoidance task in day-old chicks (white Leghorn-black Australorp). Various doses of novobiocin or menadione sodium bisulfite were administered intracranially at different times before or after training. Control chicks were administered saline at matched times. Novobiocin (650 microM) or menadione sodium bisulfite (250 microM) administered between 5.0 min pretraining and 2.5 min posttraining was found to cause a persistent loss of retention from 120 min posttraining. These data provide the first demonstration that monoADP-ribosylation is required for the maintenance of long-term memory. Furthermore, the temporal characteristics of the memory loss caused by monoADP-ribosylation inhibition appears to exclude this mechanism as a downstream effect of the well-established nitric oxide activity previously shown to occur within 40 min of passive avoidance training.     

The role of reminding in long-term memory for temporal order

Order codes are one of the three main types of information that have been hypothesized to underlie memory for the times of life events. Published evidence for the theory, however, has come exclusively from research in which brief retention intervals have been used. In the first of two studies, 101 adults judged the order of pairs of movies released 5-14 years ago, half of which shared a common major actor. There was no evidence that related films could be ordered more accurately than unrelated ones. In the second study, 88 students were presented with in-class announcements that were either related or unrelated to an earlier announcement. Three weeks later,they judged the order of the pairs of announcements. There was no difference between the accuracy for the related and the unrelated pairs. The findings do not support the proposal that the automatic creation of order information at the time of encoding contributes to autobiographical memory.     

Activation of MAPK is necessary for long-term memory consolidation following food-reward conditioning

Although an important role for the mitogen-activated protein kinase (MAPK) has been established for memory consolidation in a variety of learning paradigms, it is not known if this pathway is also involved in appetitive classical conditioning. We address this question by using a single-trial food-reward conditioning paradigm in the freshwater snail Lymnaea stagnalis. This learning paradigm induces protein synthesis-dependent long-term memory formation. Inhibition of MAPK phosphorylation blocked long-term memory consolidation without affecting the sensory and motor abilities of the snails. Thirty minutes after conditioning, levels of MAPK phosphorylation were increased in extracts from the buccal and cerebral ganglia. These ganglia are involved in the generation, modulation, and plasticity of the feeding behavior. We also detected an increase in levels of MAPK phosphorylation in the peripheral tissue around the mouth of the snails where chemoreceptors are located. Although an increase in MAPK phosphorylation was shown to be essential for food-reward conditioning, it was also detected in snails that were exposed to the conditioned stimulus (CS) or the unconditioned stimulus (US) alone, suggesting that phosphorylation of MAPK is necessary but not sufficient for learning to occur.     

In defense of experience-coding nonarbitrary temporal neural activity patterns

Various neurophysiological experiments have revealed remarkable correlations between cortical neuronal activity and subjective experiences. However, the mere presence of neuronal electrical activity does not appear to be sufficient to produce these experiences. It has been suggested that the explanation for the neural basis of consciousness might lie in understanding the reason that some types of neuronal activity possess subjective correlates and others do not. Here I propose and develop the idea that this difference may be caused by the existence of an elementary nonarbitrary linkage between temporal or spatiotemporal patterns of neuronal activity and their subjective attributes. I also show how cortical neural circuits capable of generating experience-coding patterns could emerge during evolution and brain development, due to the presence of spontaneous stochastic neuronal activity and activity-dependent synaptic plasticity. This hypothesis leads to several testable predictions, principal among which is the idea that the neural correlates of consciousness are essentially innate and universal.     

Infants' long-term memory for the sound patterns of words and voices

Infants' long-term memory for the phonological patterns of words versus the indexical properties of talkers' voices was examined in 3 experiments using the Headturn Preference Procedure (D. G. Kemler Nelson et al., 1995). Infants were familiarized with repetitions of 2 words and tested on the next day for their orientation times to 4 passages--2 of which included the familiarized words. At 7.5 months of age, infants oriented longer to passages containing familiarized words when these were produced by the original talker. At 7.5 and 10.5 months of age, infants did not recognize words in passages produced by a novel female talker. In contrast, 7.5-month-olds demonstrated word recognition in both talker conditions when presented with passages produced by both the original and the novel talker. The findings suggest that talker-specific information can prime infants' memory for words and facilitate word recognition across talkers.     

Dissociation of short- and long-term face memory: evidence from long-term recency effects in temporal lobe epilepsy

We tested whether memory deficits in temporal lobe epilepsy (TLE) are better described by a single- or dual-store memory model. To this aim, we analyzed the influence of TLE and proactive interference (PI) on immediate and 24-h long-term recency effects during face recognition in 16 healthy participants and 18 right and 21 left non-surgical TLE patients. PI in healthy participants or TLE erased the long-term recency effect, but left the immediate recency effect unaffected. Although the immediate recency effect was still visible in right TLE patients, the number of detected recency items during immediate recognition was decreased in right TLE compared to left TLE. Right TLE was also related to decreased detection of pre-recency items during delayed recognition compared to left TLE, and decreased detection of pre-recency items during immediate recognition under PI. The results show that the temporal lobes are necessary for the long-term recency effect, but not for the immediate recency effect, and thus speak for a dissociation of short- and long-term memory for faces. Right TLE is related to more severe long-term memory deficits than left TLE and is also related to additional short-term memory deficits for faces.     

Timing a week later: The role of long-term memory in temporal preparation

Temporal preparation has been investigated extensively by manipulating the foreperiod, the interval between a warning stimulus and target stimulus requiring a speeded response. Although such research has revealed many effects of both the duration and distribution of foreperiods on reaction times, the underlying cognitive mechanism is still largely unknown. Here, we test a recent proposal that temporal preparation is driven by the retrieval of memory traces of past experiences from long-term memory rather than by knowledge about upcoming events. Two groups of participants received different foreperiod distributions in an acquisition phase, which was followed a week later by a transfer phase, in which both groups received the same distribution of foreperiods. We found that the effects of the different foreperiod distributions presented in the acquisition phase were still apparent a week later during the transfer phase, as the reaction time patterns of both groups reflected the old distributions. This occurred even though both groups were provided with full information about the change in the distribution of foreperiods at the start of the transfer phase. These findings provide compelling evidence that long-term memory plays an important role in temporal preparation.     

Availability of related long-term memory during and after attention focus in working memory

In a series of four experiments, we investigated the magnitude and pattern of indirect, semantically mediated priming that emanates from various working memory (WM) processes. When such priming effects are taken to reflect temporary increases in the availability of long-term memory (ALTM), results suggest a close link between the amount and type of attention-driven processing in WM and the resulting accessibility of semantically related memory structures. Patterns of priming were equivalent when attention demands were prior to, versus concurrent with, the priming measures, suggesting that ALTM processes either require no additional cognitive resources or require resources that are independent of those underlying effortful WM processes. The results are discussed with respect to emerging evidence for long-term semantic priming and models of WM that incorporate active but unattended information as part of a limited capacity cognitive workspace.     

Spared anterograde memory for shock-probe fear conditioning after inactivation of the amygdala

Previous studies have shown that amygdala lesions impair avoidance of an electrified probe. This finding has been interpreted as indicating that amygdala lesions reduce fear. It is unclear, however, whether amygdala-lesioned rats learn that the probe is associated with shock. If the lesions prevent the formation of this association, then pretraining reversible inactivation of the amygdala should impair both acquisition and retention performance. To test this hypothesis, the amygdala was inactivated (tetrodotoxin; TTX; 1 ng/side) before a shock-probe acquisition session, and retention was tested 4 d later. The data indicated that, compared with rats infused with vehicle, rats infused with TTX received more shocks during the acquisition session, but more importantly, were not impaired on the retention test. In Experiment 2, we assessed whether the spared memory on the retention test was caused by overtraining during acquisition. We used the same procedure as in Experiment 1, with the exception that the number of shocks the rats received during the acquisition session was limited to four. Again the data indicated that amygdala inactivation did not impair performance on the retention test. These results indicate that amygdala inactivation does not prevent the formation of an association between the shock and the probe and that shock-probe deficits during acquisition likely reflect the amygdala's involvement in other processes.     

Electrodermal activity as an indicator of information processing in a nonaversive differential classical conditioning paradigm

Most investigations of information processing in human Pavlovian autonomic conditioning focussed on the role of awareness of the contingency of the conditioned stimulus (CS) and the unconditioned stimulus (UCS). While in these studies only the qualitative information of the CS (UCS occurring vs not occurring) was varied, the present experiment was designed to vary the quantitative information of the CS. In a differential conditioning paradigm, using a letter reproduction task as nonaversive UCS, electrodermal responses to different degrees of CS information content were investigated. Twenty subjects experienced two different conditions in a within subject design. In one condition only the number of letters used in the following UCS was presented as CS+, while in the other condition the CS already contained the whole UCS information. Both conditions led to significant differences in magnitudes of electrodermal responding.     

Grouping and short-term memory: Different means and patterns of grouping

Two experiments, concerned with the improving effects of grouping on auditory short-term memory, are described. In the first, temporal grouping was found to improve recall considerably, but non-temporal grouping had a much smaller effect. Temporal grouping reduced the order errors more than other errors; it also changed the pattern of the order errors. Further, it altered the shape of the serial position curve of all errors. In the second experiment, irregular patterns of temporal grouping were found to be inferior to a regular pattern. The results are discussed in terms of the time available for processing previous items during the presentation of a sequence, and the form that this processing may take.     

Topographically distinct cortical activation in episodic long-term memory: The retrieval of spatial versus verbal information

Two experiments are reported to study slow potentials in the EEG during reactivation of spatial and verbal information. Subjects had to learn associations between drawings and one, two, or three mediators (locations in Experiment 1, nouns in Experiment 2). During recall, subjects had to decide whether or not two drawings were linked to each other by a common mediator. EEG was recorded during learning and recall. Both experiments were completely equivalent. Irrespective of the quality of the mediators, response time proved to be a linear function of the numbers of mediators to be recalled. Negative slow potentials that accompanied the reactivation of information during anticipation learning and cued recall had a material-specific topography: The maximum was found over the parietal cortex for spatial information and over the left frontal cortex for verbal information. Moreover, the amplitude at these scalp locations varied with the amount of the to-be-retrieved information. The results support the claim of topographically distinct cell assemblies specialized for storage and retrieval of distinct kinds of information.     

Processing of temporal duration information in working memory after frontodorsal tumour excisions

This study aimed to test the hypothesis that impairments of temporal duration processing after frontal lobe lesions reflect deficits in executive monitoring functions rather than a domain-specific deficit in the maintenance of duration information in working memory. Patients with frontodorsal lesions, clinical controls with post-central lesions, and healthy controls performed recognition and classification tasks, which should allow for testing maintenance and monitoring functions, respectively. Results showed mild non-selective impairments of the frontal patients on both temporal and spatial recognition tasks, but a marked selective degradation on temporal classification while performance on spatial classification was unimpaired. This suggests that maintenance of duration information in working memory after frontal lesions is basically preserved but that, depending on executive task characteristics, there is a specific deficit in the strategic organization of this type of information.     

Different types of working memory binding in epilepsy patients with unilateral anterior temporal lobectomy

The medial temporal lobe is an important structure for long-term memory formation, but its role in working memory is less clear. Recent studies have shown hippocampal involvement during working memory tasks requiring binding of information. It is yet unclear whether this is limited to tasks containing spatial features. The present study contrasted three binding conditions and one single-item condition in patients with unilateral anterior temporal lobectomy.A group of 43 patients with temporal lobectomy (23 left; 20 right) and 20 matched controls were examined with a working memory task assessing spatial relational binding (object–location), non-spatial relational binding (object–object), conjunctive binding (object–colour) and working memory for single items. We varied the delay period (3 or 6 s), as there is evidence showing that delay length may modulate working memory performance.The results indicate that performance was worse for patients than for controls in both relational binding conditions, whereas patients were unimpaired in conjunctive binding. Single-item memory was found to be marginally impaired, due to a deficit on long-delay trials only.In conclusion, working memory binding deficits are found in patients with unilateral anterior temporal lobectomy. The role of the medial temporal lobe in working memory is not limited to tasks containing spatial features. Rather, it seems to be involved in relational binding, but not in conjunctive binding. The medial temporal lobe gets involved when working memory capacity does not suffice, for example when relations have to be maintained or when the delay period is long.     

Intra-medial prefrontal administration of SCH-23390 attenuates ERK phosphorylation and long-term memory for trace fear conditioning in rats

The prefrontal cortex is known to be involved in the acquisition of trace conditioning, a higher-cognitive form of Pavlovian conditioning in which a conditioned stimulus and an unconditioned stimulus are separated by a time gap. We have recently reported that medial prefrontal (mPFC) extracellular-signal regulated kinase (Erk) phosphorylation is involved in the long-term memory storage of trace fear conditioning. Because of the important role dopamine D1 receptors play in prefrontal function, such as working memory, and due to evidence that dopamine D1 receptor activity can modulate plasticity, we investigated their role in prefrontal Erk phosphorylation following trace fear conditioning. We found that inhibition of dopamine D1 receptors through intra-mPFC infusion of SCH-23390 (1 microg/0.5 microL) 15 min prior to trace fear conditioning resulted in a decrease in training-related Erk phosphorylation. Additionally, pre-training intra-mPFC infusion of SCH-23390 also resulted in the impairment of long-term retention of CS-US association. These findings implicate mPFC dopamine D1 receptor activity in the storage of long-term memory for higher-cognitive associative tasks.