The Ontogeny of Long‐Term Retention During the Second Year of Life

Developmental changes in retention were assessed in two experiments with 18‐ and 24‐month‐old infants. In both experiments, infants were tested in a deferred imitation paradigm. In Experiment 1, independent groups of infants were tested either immediately or after delays of 1, 14, 28 or 56 days. There was no age‐related difference in the spontaneous production of target actions (baseline) or in immediate imitation. There were age‐related changes in retention after longer delays. Eighteen‐month‐olds exhibited retention for 14 days and 24‐month‐olds exhibited retention for at least 56 days. In Experiment 2, the maximum duration of retention by 24‐month‐olds was assessed. Independent groups of infants were tested after 3 or 6 months. Infants exhibited some evidence of retention after 3 months; however, forgetting was complete after 6 months.     

Crawling is associated with more flexible memory retrieval by 9-month-old infants

In the present experiment, we used a deferred imitation paradigm to explore the effect of crawling on memory retrieval by 9-month-old human infants. Infants observed an experimenter demonstrate a single target action with a novel object and their ability to reproduce that action was assessed after a 24-hr delay. Some infants were tested with the demonstration stimulus in the demonstration context and some infants were tested with a different stimulus in a different context. Half of the infants in each test condition were crawling at the time of participation and half were not. Both crawling and non-crawling infants exhibited retention when tested with the demonstration stimulus in the demonstration context, but only infants who were crawling by 9 months of age exhibited retention when tested with a different stimulus in a different context. These findings demonstrate that the onset of independent locomotion is associated with more flexible memory retrieval during the first year of life.     

Contextual determinants of retrieval in three-month-old infants

Three-month old infants learned to kick to produce movement in an overhead crib mobile on a conjugate reinforcement (FRI) schedule. All infants were trained in the presence of one of two distinctive crib bumpers. In Experiment 1, infants receiving cued-recall tests in the presence of the original training bumper remembered the conditioned association 1 week later, but those tested in the presence of a different bumper did not. Two weeks later, neither training group evidenced retention of the contingency. In Experiment 2, infants who received a 3-min exposure (“a reactivation treatment”) to the original training bumper 24 h prior to the 2-week retention test exhibited excellent long-term retention, whereas those exposed to a different bumper as a reminder did not, irrepective of whether it was identical to the test bumper or not. These findings demonstrate that the context alone, even though it was never exclusively paired with reinforcement, can act as a retrieval cue.     

Olfactory context and memory retrieval in 3-month-old infants

Three experiments examineed whether olfactory stimuli can produce context-dependent effects in infant memory. In eash, 3-month-old infants learned to kick to control the movement of an overhead modile in the presence of an ambient odor. Retention was assessed 1, 3, or 5 days later. During the retention test, the olfactory context was either the same odor, a different odor, or “no odor.” At 1 day, infants exhibited retention when tested in the presence of the same odor. Infants in the no odor condition exhibited partial retention, whereas memory retrieval was completely disrupted for infants tested in the presence of the different odor. After the 3- and 5-day intervals, all groups showed forgetting. These results were not consistent with recent studies of 3-month-old infants trained and tested in the presence of a distinctive visual and auditory context. The findings suggest that olfactory stimuli are a silent feature of the environment for 3-month-old infants, but that they function as context cues in a way that differs from other sensory modalities.     

Reinstatement versus reactivation effects on active memory in infants

Reinstatement and reactivation are procedurally different reminder paradigms used with infants and children, but most developmental psychologists do not distinguish between them. In 4 experiments with 102 three-month-olds, we asked if they differ functionally as well. Independent groups of infants received either a reactivation or a reinstatement reminder 3 days after training, when the memory is active, but its specific details have been forgotten. In Experiment 1, we measured retention after increasing delays until infants forgot altogether. A single reinstatement protracted retention twice as long after training as a single reactivation. In Experiments 2-4, whether the reminder was the original training stimulus or a novel one differentially affected the duration and specificity of memory in the 2 procedures as well. These data demonstrate that the distinction between reinstatement and reactivation is not artificial. In addition to differing procedurally, reinstatement and reactivation differ functionally, with different memory-preserving effects.     

The effects of distributed training on retention of operant conditioning in human infants

Long-term retention of operant footkicking acquired in the mobile conjugate reinforcement paradigm was assessed as a function of the distribution of training time. In the first study, 3-month-old infants were trained for either one 18-min session or for two 9-min or three 6-min sessions separated by 24-hr intervals. All infants exhibited retention during a test administered immediately after training, but only those trained in a single session continued to perform the conditioned response during cued-recall tests 7 or 14 days later. Infants trained in three sessions showed no evidence of remembering the contingency even after a week. A warm-up decrement, seen in the day-to-day performance of infants in the distributed conditions, was eliminated in the second study by the interpolation of a nonreinforcement period at the outset of daily sessions. This procedure also enhanced long-term retention such that infants trained in three 6-min session now remembered the contingency for 14 days and did not differ from infants who had received a single 18-min session. Whether distributed training facilitates or impairs long-term retention appears to depend on the opportunity for infants to acquire a sufficient number (or kind) of effective retrieval cues during original learning.     

The effect of a salient odor context on memory retrieval in young infants

Three-month-old infants were trained to move a mobile in the presence of a coconut or cherry odor (context). One or 5 days later, the infants were tested for retrieval in the presence of either the same odor, the alternate odor, or no odor. Infants tested with the training odor displayed retention at both intervals; retention was not seen at either interval in the alternate odor or no odor conditions. These data suggest that the odor combines with the mobile to form a compound-stimulus representation of the learned task whose presence after both short (1 day) and long (5 days) intervals is a necessary retrieval cue.     

Inhibition of mTOR by rapamycin in the amygdala or hippocampus impairs formation and reconsolidation of inhibitory avoidance memory

Mammalian target of rapamycin (mTOR), a central regulator of protein synthesis in neurons, has been implicated in synaptic plasticity and memory. Here we show that mTOR inhibition by rapamycin in the basolateral amygdala (BLA) or dorsal hippocampus (DH) impairs both formation and reconsolidation of memory for inhibitory avoidance (IA) in rats. Male Wistar rats received bilateral infusions of vehicle or rapamycin into the BLA or DH before or after IA training or retrieval. Memory retention was tested at different time points after drug infusion. Rapamycin impaired long-term IA retention when given before or immediately after training or retrieval into the BLA. When infused into the DH, rapamycin produced memory impairment when given before training or immediately after retrieval. The impairing effects of post-retrieval rapamycin required memory retrieval and were not reversed by a reminder shock. The results provide the first evidence that mTOR in the BLA and DH might play a role in IA memory reconsolidation.     

Reinstatement Effects on Retention at 3 Months of Age

In three experiments with 56 3-month-olds, we examined the effect of different numbers of reinstatements (reminders) on long-term retention. Infants learned to move a crib mobile by kicking and subsequently received one, two, or three reinstatements. Each reinstatement was a partial training episode one-sixth the duration of original training. Presenting a single reinstatement when the memory was inactive failed to recover it 1 day later (Experiment 1), but increasing the number of reinstatements when the memory was active to two (Experiment 2) and three (Experiment 3) progressively protracted retention. Although 3-month-olds typically remember for less than 6 days, after three reinstatements they still exhibited retention 6 weeks after training. Untrained controls who received an identical regimen of reinstatements exhibited no retention. These results demonstrate that periodic reinstatements can maintain young infants' retention over long delays and that the state of the memory at the time of reinstatement is critical to its effectiveness.     

Motion tracking of parents’ infant‐ versus adult‐directed actions reveals general and action‐specific modulations

Parents tend to modulate their movements when demonstrating actions to their infants. Thus far, these modulations have primarily been quantified by human raters and for entire interactions, thereby possibly overlooking the intricacy of such demonstrations. Using optical motion tracking, the precise modulations of parents’ infant‐directed actions were quantified and compared to adult‐directed actions and between action types. Parents demonstrated four novel objects to their 14‐month‐old infants and adult confederates. Each object required a specific action to produce a unique effect (e.g. rattling). Parents were asked to demonstrate an object at least once before passing it to their demonstration partner, and they were subsequently free to exchange the object as often as desired. Infants’ success at producing the objects’ action‐effects was coded during the demonstration session and their memory of the action‐effects was tested after a several‐minute delay. Indicating general modulations across actions, parents repeated demonstrations more often, performed the actions in closer proximity and demonstrated action‐effects for longer when interacting with their infant compared to the adults. Meanwhile, modulations of movement size and velocity were specific to certain action‐effect pairs. Furthermore, a ‘just right’ modulation of proximity was detected, since infants’ learning, memory, and parents’ prior evaluations of their infants’ motor abilities, were related to demonstrations that were performed neither too far from nor too close to the infants. Together, these findings indicate that infant‐directed action modulations are not solely overall exaggerations but are dependent upon the characteristics of the to‐be learned actions, their effects, and the infant learners.     

Memory by association: Integrating memories prolongs retention by two-year-olds

Recalling one memory often leads to the recollection of other memories that share overlapping features. This phenomenon, spreading activation, was originally documented in studies conducted with verbal adults, and more recently, it has been demonstrated with preverbal infants. Here, we examine the effect of spreading activation on long-term retention by 2-year-olds. Participants were tested in the Visual Recognition Memory (VRM) paradigm and the deferred imitation paradigm. Typically, infants of this age exhibit retention in the VRM paradigm for 24 h, while they exhibit retention in the deferred imitation paradigm for at least 8 weeks. In the present experiment, we paired these tasks together during original encoding and tested infants after an 8-week delay. Two-year-olds exhibited retention in both tasks. That is, when these two tasks initially occurred together – one task that is extremely memorable and one that is not – retrieving the memory of the more memorable task cued retrieval of the less memorable task, extending its longevity.     

The effects of the dopamine D1 receptor antagonist SCH23390 on memory reconsolidation following reminder-activated retrieval in day-old chicks

This series of experiments examined the involvement of the dopamine D1 receptor antagonist, SCH23390, on memory reconsolidation following reminder-activated retrieval. Day-old male New HampshirexWhite Leghorn chicks were trained on a single trial passive avoidance task. A dose of 0.5 mg/kg of SCH23390 was administered subcutaneously 5 min before reminder trials, which were presented at 30, 60, and 90 min following training. Memory deficits were observed when reminder trials were presented at 30 and 60 min following training, but not when a reminder was presented at 90 min. No effect on memory retention was observed when reminder trials were not presented, suggesting that reconsolidation mechanisms were both contingent on the presentation of the reminder and independent of the consolidation process. Following a reminder presented at 60 min post-training, deficits in memory retention emerged between 45 and 60 min. The deficit was prolonged, lasting for up until 48 h after reminder presentation. The results indicate an important role for the D1 receptor in reconsolidation processes.     

Verbal cues facilitate memory retrieval during infancy

In three experiments, 18-month-olds were tested in a deferred imitation paradigm. Some infants received verbal information during the demonstration and at the time of the test (full narration), and some did not (empty narration). When tested after a 4-week delay, infants given full narration exhibited superior retention relative to infants given empty narration (Experiment 1). This retention advantage appears to be due to the effects of verbal cues at the time of memory retrieval. There was no effect of verbal cues that were presented only at the time of original encoding (Experiment 2A). Furthermore, infants who received verbal cues only at the time of retrieval exhibited superior retention relative to infants who received verbal cues only at the time of original encoding (Experiment 2B). These findings demonstrate that verbal cues can enhance memory retrieval by participants who are not yet fluent speakers themselves.     

Post-training glucose administration attenuates forgetting of passive-avoidance conditioning in 18-day-old rats.

The study of memory modulation in infant rats has typically focused on reminder/retrieval treatments involving reexposure to components of the internal or external training context. Rarely have studies employed pharmacological treatments to investigate the neurochemical substrates of memory storage in preweanling rats. The present study investigated the effect of 100 mg/kg of glucose, a common memory modulator in adult mammals, on memory for passive-avoidance conditioning in 18-day-old Sprague-Dawley rats. Subjects that were administered an immediate post-training injection of glucose performed significantly better, on a retention test 24 h following training, than those animals that received saline. The glucose group also performed comparably to a control group that was tested 10 min following training. These results are consistent with those of the memory modulation literature in adults and suggest that the rapid rate of forgetting in immature organisms may be the result of a deficiency in a general memory modulatory system.     

A systemically administered beta-adrenoceptor antagonist blocks corticosterone-induced impairment of contextual memory retrieval in rats

Several studies have reported that glucocorticoids impair memory retrieval. The present study examined in male Sprague-Dawley rats the effects of systemically administered corticosterone on retrieval of memory for inhibitory avoidance training. Corticosterone (3.0mg/kg, s.c.) injected 30min before retention testing, 48h after training, significantly impaired retention performance, as compared to vehicle treatment, of rats tested in the training context. In contrast, corticosterone administration did not impair retrieval when rats were tested for retention in a different context. Corticosterone did also not impair retention performance of rats given a mild-intensity footshock that resulted in only weak, non-contextual memory. These findings strongly suggest that corticosterone selectively impaired retrieval of contextual information associated with the training context. The centrally acting beta-adrenoceptor antagonist propranolol (2.0mg/kg), co-administered in a dose that did not affect retention performance alone, blocked the impairment in contextual memory retrieval induced by corticosterone. These findings provide evidence for the view that glucocorticoids interact with noradrenergic mechanisms in influencing memory retrieval.     

Contextual coding and recoding of infants' memories.

Six-month-old infants recognize a well-learned cue 24 h after training in the original context but not in a different one. In the present studies, we demonstrated that this contextually based retrieval deficit could be overcome if, following some training experience in one context, infants were briefly and passively exposed to a novel context. The impact of this procedure was retroactive, leading the immediately preceding training episode to be contextually recoded. However, it did not override the debilitating effect of a novel context on memory reactivation 3 weeks later and, surprisingly, it blocked the effectiveness of the actual training context as a reminder. We conclude that each individual training episode is encoded in terms of the context in which it occurs. The memory of a given episode, however, is recoded in terms of the most recent context in which it is active. This mechanism permits memories that are highly context-specific to be updated and subsequently retrieved in new and potentially appropriate settings.     

Substituting new details for old? Effects of delaying postevent information on infant memory

How that which we remember is selectively distorted by new information was studied in 3-monthold infants who learned to move a particular crib mobile by operant foot kicking. Infants who were passively exposed to a novel mobile 1, 2, or 3 days later subsequently treated the novel mobile as if they had actually been trained with it. Also, after the longest exposure delay, they no longer recognized the original mobile. Likewise, when the novel mobile was exposed after the longest delay, it could prime the forgotten training memory in a reactivation paradigm, but the original mobile no longer could. These data reveal that what we remember about an event is selectively distorted by what we encounter later. Moreover, the later in the retention interval we encounter new postevent information, the greater is its impact on retention.     

The reciprocal relation between sleep and memory in infancy: Memory‐dependent adjustment of sleep spindles and spindle‐dependent improvement of memories

Sleep spindle activity in infants supports their formation of generalized memories during sleep, indicating that specific sleep processes affect the consolidation of memories early in life. Characteristics of sleep spindles depend on the infant's developmental state and are known to be associated with trait‐like factors such as intelligence. It is, however, largely unknown which state‐like factors affect sleep spindles in infancy. By varying infants’ wake experience in a within‐subject design, here we provide evidence for a learning‐ and memory‐dependent modulation of infant spindle activity. In a lexical‐semantic learning session before a nap, 14‐ to 16‐month‐old infants were exposed to unknown words as labels for exemplars of unknown object categories. In a memory test on the next day, generalization to novel category exemplars was tested. In a nonlearning control session preceding a nap on another day, the same infants heard known words as labels for exemplars of already known categories. Central–parietal fast sleep spindles increased after the encoding of unknown object–word pairings compared to known pairings, evidencing that an infant's spindle activity varies depending on its prior knowledge for newly encoded information. Correlations suggest that enhanced spindle activity was particularly triggered, when similar unknown pairings were not generalized immediately during encoding. The spindle increase triggered by previously not generalized object–word pairings, moreover, boosted the formation of generalized memories for these pairings. Overall, the results provide first evidence for a fine‐tuned regulation of infant sleep quality according to current consolidation requirements, which improves the infant long‐term memory for new experiences.     

Memories in drosophila heat-box learning

Learning and memory processes of operant conditioning in the heat-box are analyzed. In a search for conditioning parameters leading to high retention scores, intermittent training is shown to give better results than those of continuous training. Immediate retention tests contain two memory components, a spatial preference for one side of the chamber and a “stay-where-you-are-effect.” Intermittent training strengthens the latter. In the second part, memory dynamics is investigated. Flies are trained in one chamber and tested in a second one after a brief reminder training. With this direct transfer, memory scores reflect an associative learning process in the first chamber. To investigate memory retention after extended time periods, indirect transfer experiments are performed. The fly is transferred to a different environment between training and test phases. With this procedure, an aftereffect of the training can still be observed 2 h later. Surprisingly, exposure to the chamber without conditioning also leads to a memory effect in the indirect transfer experiment. This exposure effect reveals a dispositional change that facilitates operant learning during the reminder training. The various memory effects are independent of the mushroom bodies.     

Who's my little monkey? Effects of infant‐directed speech on visual retention in infant rhesus macaques

Both human and nonhuman primate adults use infant‐directed facial and vocal expressions across many contexts when interacting with infants (e.g., feeding, playing). This infant‐oriented style of communication, known as infant‐directed speech (IDS), seems to benefit human infants in numerous ways, including facilitating language acquisition. Given the variety of contexts in which adults use IDS, we hypothesized that IDS supports learning beyond the linguistic domain and that these benefits may extend to nonhuman primates. We exposed 2.5‐month‐old rhesus macaque infants (N = 15) to IDS, adult‐directed speech (ADS), and a non‐social control (CTR) during a video presentation of unrelated stimuli. After a 5‐ or 60‐minute delay, infants were shown the familiar video side‐by‐side with a novel video. Infants exhibited a novelty preference after the 5‐minute delay, but not after the 60‐minute delay, in the ADS and CTR conditions, and a novelty preference in the IDS condition only after the 60‐minute delay. These results are the first to suggest that exposure to IDS affects infants’ long‐term memory, even in non‐linguistic animals.