The procerebrum is necessary for odor-aversion learning in the terrestrial slug Limax valentianus

The terrestrial slug Limax has a highly developed ability to associate the odor of some foods (e.g., carrot juice) with aversive stimuli such as the bitter taste of quinidine solution. The procerebrum (PC) is a part of the slug's brain thought to be involved in odor-aversion learning, but direct evidence is still lacking. Here we present evidence showing that the PC is essential for odor-aversion learning. Unlike sham-operated slugs, PC ablation 7 d prior to conditioning showed that most slugs did not avoid carrot juice in the memory retention test conducted 24 h after the conditioning. Slugs with the PC ablated 3 h, 1 d, 3 d, or 7 d after conditioning and examined by the memory retention test at 3 d after the PC ablation were also less likely to avoid carrot juice than sham-operated slugs. The PC ablation did not damage the ability of the slugs to sense attractive odor (everyday food) or innately aversive odor (onion or garlic). These results demonstrate that the PC is a necessary component in the retention and/or retrieval of odor-aversion memory.     

Unilateral memory storage in the procerebrum of the terrestrial slug Limax

In most sensory modalities, neuronal inputs are bilaterally processed in a higher center. In some animal species, however, functional lateralization is sometimes observed in the sensory processing at the higher level. For the terrestrial slug Limax, olfaction is the most important sensory modality and this slug can acquire odor-aversion memories. Previously, it has been demonstrated in bilateral PC ablation experiments that the procerebrum (PC) is necessary for odor-aversion memory, and that the PC is the memory storage site. On the other hand, it has been hypothesized that only the unilateral PC is used for odor-aversion learning. Here we demonstrated that the number of the slugs with intact memory performance was reduced by approximately 50% when the PC was surgically ablated only unilaterally before or after conditioning. There was no difference in the memory performance of the right vs. the left PC-ablated slugs. However, memory deficit from unilateral PC ablation was not observed when the ipsilateral tentacles were also amputated at the same time. We also showed that there was no lateral memory transfer from one PC to the other, after up to 7 days post-conditioning. Our results demonstrated clearly that either the left or right PC is randomly used for olfactory learning, and that the side of use is determined at the level of the olfactory ascending pathway to the PC.     

Reactivation-dependent changes in memory states in the terrestrial slug Limax flavus

The change in memory state in the terrestrial slug Limax flavus was studied using cooling-induced retrograde amnesia. Slugs were first conditioned to avoid carrot odor and then a second conditioning procedure was applied 1, 3, 6, or 7 days after the first conditioning trial. Cooling the slugs to approximately 1 degrees C on day 7 immediately after the presentation of the odor used in the conditioning resulted in retrograde amnesia in the slugs that were subject to a second conditioning on day 6 or 7, but not in slugs that were subject to a second conditioning on day 1 or 3. Next, second-order conditioning was used as the second conditioning procedure to distinguish the memory acquired in the first conditioning from that acquired in the second conditioning and similar results were obtained. These results suggest that the reactivation of memory altered the memory state from a cooling-insensitive state to a cooling-sensitive one. A possible model for memory states is discussed.     

Nitric oxide is involved in appetitive but not aversive olfactory learning in the land mollusk Limax valentianus

The land slug Limax performs both aversive and appetitive olfactory learning, and we investigated neurotransmitters involved in each type of learning. Slugs were conditioned by presenting a vegetable juice (appetitive conditioning) or a mixture of vegetable juice and quinidine (aversive conditioning), and the latency to reach the juice became shorter (appetitive conditioning) or longer (aversive conditioning) after conditioning. L-NAME injected either before conditioning or testing blocked the reduction in latency in appetitive conditioning but had no significant effects in aversive conditioning. 5,7-dihydroxytryptamine had no significant effects in appetitive conditioning. These results suggest different mechanisms for appetitive and aversive learning.     

Deficits in verbal long-term memory and learning in children with poor phonological short-term memory skills

Possible links between phonological short-term memory and both longer term memory and learning in 8-year-old children were investigated in this study. Performance on a range of tests of long-term memory and learning was compared for a group of 16 children with poor phonological short-term memory skills and a comparison group of children of the same age with matched nonverbal reasoning abilities but memory scores in the average range. The low-phonological-memory group were impaired on longer term memory and learning tasks that taxed memory for arbitrary verbal material such as names and nonwords. However, the two groups performed at comparable levels on tasks requiring the retention of visuo-spatial information and of meaningful material and at carrying out prospective memory tasks in which the children were asked to carry out actions at a future point in time. The results are consistent with the view that poor short-term memory function impairs the longer term retention and ease of learning of novel verbal material.     

Deficits in verbal long-term memory and learning in children with poor phonological short-term memory skills

Possible links between phonological short-term memory and both longer term memory and learning in 8-year-old children were investigated in this study. Performance on a range of tests of long-term memory and learning was compared for a group of 16 children with poor phonological short-term memory skills and a comparison group of children of the same age with matched nonverbal reasoning abilities but memory scores in the average range. The low-phonological-memory group were impaired on longer term memory and learning tasks that taxed memory for arbitrary verbal material such as names and nonwords. However, the two groups performed at comparable levels on tasks requiring the retention of visuo-spatial information and of meaningful material and at carrying out prospective memory tasks in which the children were asked to carry out actions at a future point in time. The results are consistent with the view that poor short-term memory function impairs the longer term retention and ease of learning of novel verbal material.     

Factors affecting the learning of foreign language vocabulary: Imagery keyword mediators and phonological short-term memory

In order to investigate the cognitive processes involved in learning Foreign Language (FL) vocabulary, this study evaluates different methods of instruction. It demonstrates that keyword techniques are effective for receptive learning but that repetition is a superior strategy for learning to produce the foreign word. Performance is optimal when learners combine both strategies. The nature of the keyword is crucial--whereas imageable noun keywords promote learning, verb keywords actually impede it. A theoretical analysis of the roles of phonological short-term memory, imagery, and lexical factors in FL vocabulary learning is presented.     

Acute ethanol ingestion impairs appetitive olfactory learning and odor discrimination in the honey bee

Invertebrates are valuable models for increasing our understanding of the effects of ethanol on the nervous system, but most studies on invertebrates and ethanol have focused on the effects of ethanol on locomotor behavior. In this work we investigate the influence of an acute dose of ethanol on appetitive olfactory learning in the honey bee (Apis mellifera), a model system for learning and memory. Adult worker honey bees were fed a range of doses (2.5%, 5%, 10%, or 25%) of ethanol and then conditioned to associate an odor with a sucrose reward using either a simple or differential conditioning paradigm. Consumption of ethanol before conditioning significantly reduced both the rate of acquisition and the asymptotic strength of the association. Honey bees also exhibited a dose dependent reduction in arousal/attention during conditioning. Consumption of ethanol after conditioning did not affect recall 24h later. The observed deficits in acquisition were not due to the affect of ethanol on gustatory sensitivity or motor function. However, honey bees given higher doses of ethanol had difficulty discriminating amongst different odors suggesting that ethanol consumption influences olfactory processing. Taken together, these results demonstrate that an acute dose of ethanol affects appetitive learning and olfactory perception in the honey bee.     

The effects of word co-occurrence on short-term memory: associative links in long-term memory affect short-term memory performance

In immediate serial recall tasks, high-frequency words are recalled better than low-frequency words. This has been attributed to high-frequency words' being better represented and providing more effective support to a redintegration process at retrieval (C. Hulme et al., 1997). In studies of free recall, there is evidence that frequency of word co-occurrence, rather than word frequency per se, may explain the recall advantage enjoyed by high-frequency words (J. Deese, 1960). The authors present evidence that preexposing pairs of low-frequency words, so as to create associative links between them, has substantial beneficial effects on immediate serial recall performance. These benefits, which are not attributable to simple familiarization with the words per se, do not occur for high-frequency words. These findings indicate that associative links between items in long-term memory have important effects on short-term memory performance and suggest that the effects of word frequency in short-term memory tasks are related to differences in interitem associations in long-term memory.     

Behavioral modulation induced by food odor aversive conditioning and its influence on the olfactory responses of an oscillatory brain network in the slug Limax marginatus

We compared behaviorally and physiologically the olfactory responses of slugs (Limax marginatus) that had been subjected to aversive, appetitive, or unpaired training with food odors (carrot or cucumber). In the aversive training, the slugs were exposed to the food odor as a conditioned stimulus (CS), and then quinidine sulfate solution as an unconditioned stimulus (UCS) was immediately applied to the lip of the slugs. This training caused a decrease in preference level for the CS. The unpaired training, in which the CS and the UCS were presented to the slugs with a 5-min interval, induced no change in the preference level for the CS. In the appetitive training, the slugs were allowed to eat the CS odor source without UCS application. When we used nonstarved slugs, it was found that the preference level for the CS increased upon the appetitive training. These results indicate that each training changed the preference for the odors in a characteristic manner. In the physiological experiments, we used brain-inferior tentacular nose preparations isolated from slugs and investigated the olfactory responses of the oscillations in the local field potential (LFP) of the procerebral (PC) lobe. We found that odor presentation induced various types of changes in the LFP oscillation frequency, although the rate of occurrence of the frequency modulation differed between odors used in the aversive and the unpaired training (aversive-conditioned and unpaired odors). The aversive-conditioned odors induced a decrease in the oscillatory frequency. Unpaired odors did not change it. Moreover, odors used in the appetitive training (appetitive-conditioned odors) induced an increase in the frequency. Thus, it was considered that those modulations of PC lobe oscillatory activity were independent of odor and reflected learned preference for odors.     

Contrasting contributions of phonological short-term memory and long-term knowledge to vocabulary learning in a foreign language

The contributions of phonological short-term memory and existing foreign vocabulary knowledge to the learning of new words in a second language were compared in a sample of 40 Greek children studying English at school. The children's speed of learning new English words in a paired-associate learning task was strongly influenced by their current English vocabulary, but was independent of phonological memory skill, indexed by nonword repetition ability. However, phonological memory performance was closely linked to English vocabulary scores. The findings suggest that in learners with considerable familiarity with a second language, foreign vocabulary acquisition is mediated largely by use of existing knowledge representations.     

Inactivation of the basolateral amygdala impairs the retrieval of recent and remote taste-potentiated odor aversion memory

Memory reorganization as a time-dependent process can be investigated using various learning tasks such as the taste-potentiated odor aversion (TPOA). In this paradigm rats acquire a strong aversion to an olfactory cue presented simultaneously with a gustatory cue. Together these cues are paired with a delayed visceral illness. The basolateral amygdaloid nucleus (BLA) plays a key role in TPOA acquisition but its involvement in retrieval remains unclear. We investigated the involvement of the BLA in either recent or remote retrieval of TPOA. In each case, the number of licks observed in response to the presentation of either the odor or the taste was used to assess retrieval. Before the retrieval test, rats received a bilateral infusion of lidocaine to inactivate the BLA. We observed that both recent and remote TPOA retrieval tests induced by the odor presentation were disrupted in the lidocaine-injected rats. By contrast, the BLA inactivation had no effect upon the aversion towards the taste cue regardless of the time of retrieval. The present study provides evidence that BLA functioning is necessary for retrieval of aversive odor memory, even with a long post-acquisition delay.     

Reconsolidation of a context long-term memory in the terrestrial snail requires protein synthesis

We investigated the influence of the protein synthesis blocker anisomycin on contextual memory in the terrestrial snail Helix. Prior to the training session, the behavioral responses in two contexts were similar. Two days after a session of electric shocks (5 d) in one context only, the context conditioning was observed as the significant difference of behavioral response amplitudes in two contexts. On the day following testing of context learning, a session of "reminding" was performed, immediately after which the snails were injected with anisomycin or vehicle. Testing of long-term context memory has shown that only anisomycin injections impaired the context conditioning. In control series, the snails were injected after the training session with anisomycin/saline without reminding, and no impairment of the long-term context memory was observed, while injection of anisomycin during the training session completely abolished the long-term memory. No effects of anisomycin on the short-term memory were observed. Surprisingly, injection of anisomycin after the reminding combined with reinforcing stimuli elicited no effect on the context memory. Differences between single-trial and multisession learning are discussed.     

Facilitative effects of forgetting from short-term memory on growth of long-term memory in retardates

Competing explanations of the beneficial effect of spacing in retardate discrimination learning were tested. Three-trial multidimensional (“junk”) problems were presented concurrently to educable retardates with MAs over 8 years. The spacing interval spearating Trials 1 and 2 was filled with either zero, four, or eight interpolations, the interpolations being either trials on similar junk problems or repetitions of a single, well-learned dot-pattern discrimination. The principal findings were that: (1) while short-term retention of junk problems (as measured by Trial 2 performance) suffered greater interference from similar junk interpolations than from dot-pattern interpolations, long-term learning (as measured by a delayed Trial 3) was superior following the highly interfering junk problems; and (2) spacing facilitated learning only following highly interfering interpolations. These results are inconsistent with consolidation and rehearsal theories but support the prediction of the Sperber. Greenfield, and House (Journal of Experimental Psychology, 1973, 99) spacing model that forgetting from short-term memory facilitates retardate learning.     

Task-irrelevant distractors in the delay period interfere selectively with visual short-term memory for spatial locations

Visual short-term memory (VSTM) enables the representation of information in a readily accessible state. VSTM is typically conceptualized as a form of “active” storage that is resistant to interference or disruption, yet several recent studies have shown that under some circumstances task-irrelevant distractors may indeed disrupt performance. Here, we investigated how task-irrelevant visual distractors affected VSTM by asking whether distractors induce a general loss of remembered information or selectively interfere with memory representations. In a VSTM task, participants recalled the spatial location of a target visual stimulus after a delay in which distractors were presented on 75% of trials. Notably, the distractor’s eccentricity always matched the eccentricity of the target, while in the critical conditions the distractor’s angular position was shifted either clockwise or counterclockwise relative to the target. We then computed estimates of recall error for both eccentricity and polar angle. A general interference model would predict an effect of distractors on both polar angle and eccentricity errors, while a selective interference model would predict effects of distractors on angle but not on eccentricity errors. Results showed that for stimulus angle there was an increase in the magnitude and variability of recall errors. However, distractors had no effect on estimates of stimulus eccentricity. Our results suggest that distractors selectively interfere with VSTM for spatial locations.     

Some relevant factors in the transfer of material from short-term to long-term memory

The relationship between short-term memory (STM) and long-term memory (LTM) for digits was investigated by means of a Hebb-type experiment, viz. a presentation of a series of nine-digit numbers, in which a certain number recurs at intervals. Improvement in performance, with regard to the critical, or recurring, number was found when the rehearsal grouping was three-three-and-three, but was absent when there were no opportunities for rehearsal or when the rehearsal strategy was grouping five-and-four or searching for systematic numbers.

However, if an overt recall response was given on each occasion the recurring number was presented, improved performance was found even with rehearsal stratgegy five-and-four.

The conclusion was drawn that rehearsal is the main transferring mechanism from STM to LTM, with the occurrence of an over recall response as a subsidiary factor.     

The amygdala is involved in the modulation of long-term memory, but not in working or short-term memory

Rats with cannulae implanted in the junction between the central and the basolateral nuclei of the amygdala were trained in one-trial step-down inhibitory avoidance and tested at 3 s for working memory (WM) or 1.5 or 24 h later for short-term memory (STM) and long-term memory (LTM), respectively. Several drugs were infused 6 min prior to training in the animals in which WM was measured or 0 min posttraining in those in which STM and LTM were measured: the glutamate receptor antagonists CNQX (0.5 microg) and AP5 (5.0 microg), the indirect GABA A receptor antagonist picrotoxin (0.08 microg), the cholinergic muscarinic receptor blocker scopolamine (2. 0 microg), norepinephrine (0.3 microg), the protein kinase C inhibitor staurosporin (1.0 microg), or the calcium/calmodulin dependent protein kinase II inhibitor Kn-62 (3.5 ng). None of the drugs had any effect on either WM or STM. All had, as previously shown, strong effects on LTM: picrotoxin and norepinephrine enhanced it, and CNQX, AP5, scopolamine, Kn-62, and staurosporin inhibited it. The results do not support the idea that memory of this task is formed in the amygdala; they indicate that the amygdala is not involved in WM or STM processing and support the idea that the amygdala modulates LTM storage processes carried out elsewhere.