The Isolation, Primacy, and Recency Effects Predicted by an Adaptive LTD/LTP Threshold in Postsynaptic Cells

An item that stands out (is isolated) from its context is better remembered than an item consistent with the context. This isolation effect cannot be accounted for by increased attention, because it occurs when the isolated item is presented as the first item, or by impoverished memory of nonisolated items, because the isolated item is better remembered than a control list consisting of equally different items. The isolation effect is seldom experimentally or theoretically related to the primacy or the recency effects—that is, the improved performance on the first few and last items, respectively, on the serial position curve. The primacy effect cannot easily be accounted for by rehearsal in short-term memory because it occurs when rehearsal is eliminated. This article suggests that the primacy, the recency, and the isolation effects can be accounted for by experience-dependent synaptic plasticity in neural cells. Neurological empirical data suggest that the threshold that determines whether cells will show long-term potentiation (LTP) or long-term depression (LTD) varies as a function of recent postsynaptic activity and that synaptic plasticity is bounded. By implementing an adaptive LTP-LTD threshold in an artificial neural network, the various aspects of the isolation, the primacy, and the recency effects are accounted for, whereas none of these phenomena are accounted for if the threshold is constant. This theory suggests a possible link between the cognitive and the neurological levels.     

Understanding priming of color-singleton search: Roles of attention at encoding and “retrieval”

We investigated how the performance of a color-singleton search (the search for a single odd-colored item among homogeneously colored distractors) left a persistent memory trace (lasting up to six intervening trials or ～17 sec) that facilitated a subsequent color-singleton search (when the same targetdistractor color combination was repeated). Specifically, we investigated the roles of attention in the encoding and “retrieval” stages of this priming effect by intermixing trials in which the target location was precued by an onset cue. We found that the encoding of both target and distractor colors was automatic in that whether or not observers had to use color in locating the target in the preceding trial did not substantially affect priming. However, priming required that the color-singleton item be attended in the preceding trial. Once a color singleton display was encoded, our results indicated that priming facilitated the direction of attention to the color-singleton target on a subsequent trial. In short, when a color-singleton item happened to be a critical item to be attended in one situation, another color-singleton item defined by the same color combination tended to attract attention in subsequent encounters.     

Understanding priming of color-singleton search: roles of attention at encoding and "retrieval"

We investigated how the performance of a color-singleton search (the search for a single odd-colored item among homogeneously colored distractors) left a persistent memory trace (lasting up to six intervening trials or approximately 17 sec) that facilitated a subsequent color-singleton search (when the same target-distractor color combination was repeated). Specifically, we investigated the roles of attention in the encoding and "retrieval" stages of this priming effect by intermixing trials in which the target location was precued by an onset cue. We found that the encoding of both target and distractor colors was automatic in that whether or not observers had to use color in locating the target in the preceding trial did not substantially affect priming. However, priming required that the color-singleton item be attended in the preceding trial. Once a color singleton display was encoded, our results indicated that priming facilitated the direction of attention to the color-singleton target on a subsequent trial. In short, when a color-singleton item happened to be a critical item to be attended in one situation, another color-singleton item defined by the same color combination tended to attract attention in subsequent encounters.     

Prospective remembering: Perceptually driven or conceptually driven processes?

Converging experimental operations and several prospective memory tasks were used across three experiments to determine the extent to which prospective remembering is supported by datadriven versus conceptually driven processes. In all experiments, subjects were asked to perform an action when a target item later occurred. When the semantic context changed from encoding to test, prospective memory significantly declined (Experiment 1). When the target event (the item, which in its subsequent appearance in the experiment was the signal to perform the action) was presented as a word (relative to picture presentation, Experiment 2) or was encoded nonsemantically (relative to semantic encoding, Experiment 3), there was a decline in prospective memory performance. Dividing attention during prospective memory retrieval substantially reduced prospective memory performance (Experiment 3). The results of this research indicated that prospective memory is largely conceptually driven, and it behaves more similarly to direct rather than indirect conceptual tests. We suggest that prospective remembering of the type studied here is mediated by a reflexive episodic associative memory system as proposed by Moscovitch (1994).     

Distributing versus focusing attention in visual short-term memory

Visual short-term memory (VSTM) is traditionally considered a robust form of visual memory resistant to interference from subsequent visual input. This study shows that the robustness of VSTM depends on the way attention is allocated in VSTM. When attention is distributed across multiple memory items, VSTM for these items is vulnerable to interference from subsequent input, including passively viewed images and the postchange testing displays. Yet attention can readjust its focus in VSTM on the basis of an attentional orienting cue presented long after encoding. When attention is oriented to a particular memorized item, the memory is resistant to subsequent interference. This effect, however, is eliminated when the subset of items demanding focal attention exceeds one, suggesting that orienting attention in VSTM is less flexible than orienting attention in perception. We propose that the robustness of VSTM is influenced by whether attention is focused or distributed in VSTM.     

Neural evidence for the contribution of holistic processing but not attention allocation to the other-race effect on face memory

Multiple mechanisms have been suggested to contribute to the other-race effect on face memory, the phenomenon of better memory performance for own-race than other-race faces. Here, two of these mechanisms, increased attention allocation and greater holistic processing during memory encoding for own-race than other-race faces, were tested in two separate experiments. In these experiments event-related potentials were measured during study (the difference due to memory, Dm) and test phase (old/new effects) to examine brain activation related to memory encoding and retrieval, allowing for selective investigations of these memory sub-processes. In Experiment 1, participants studied own-race (Caucasian) and other-race (Chinese) faces under focused or divided attention. In Experiment 2, participants studied own-race (Caucasian) and other-race (African American) faces presented upright or upside down (i.e., inverted). Both experiments showed decreases in memory performance when attention allocation or holistic processing was reduced, but these effects were similar for own-race and other-race faces. Manipulations of holistic processing, but not attention allocation, influenced the neural other-race effects during memory encoding. Inverted own-race faces showed similar neural patterns as upright other-race faces, indicating that when holistic processing of own-race faces was reduced, these faces were encoded similarly as upright other-race faces. No influences of the experimental manipulations on other-race effects during memory retrieval were found. The present study provides the first neural evidence that increased holistic processing during memory encoding contributes to the other-race effect on face memory.     

Monkey memory: same/different concept learning, serial probe acquisition, and probe delay effects

Three rhesus monkeys were trained and tested in a same/different task with six successive sets of 70 item pairs to an 88% accuracy on each set. Their poor initial transfer performance (55% correct) with novel stimuli improved dramatically to 85% correct following daily item changes in the training stimuli. They acquired a serial-probe-recognition (SPR) task with variable (1-6) item list lengths. This SPR acquisition, although gradual, was more rapid for the monkeys than for pigeons similarly trained. Testing with a fixed list length of four items at different delays between the last list item and the probe test item revealed changes in the serial-position function: a recency effect (last items remembered well) for 0-s delay, recency and primacy effects (first and last list items remembered well) for 1-, 2-, and 10-s delays, and only a primacy effect for the longest 30-s delay. These results are compared with similar ones from pigeons and are discussed in relation to theories of memory processing.     

Where perception meets memory: A review of repetition priming in visual search tasks

What we have recently seen and attended to strongly influences how we subsequently allocate visual attention. A clear example is how repeated presentation of an object’s features or location in visual search tasks facilitates subsequent detection or identification of that item, a phenomenon known as priming. Here, we review a large body of results from priming studies that suggest that a short-term implicit memory system guides our attention to recently viewed items. The nature of this memory system and the processing level at which visual priming occurs are still debated. Priming might be due to activity modulations of low-level areas coding simple stimulus characteristics or to higher level episodic memory representations of whole objects or visual scenes. Indeed, recent evidence indicates that only minor changes to the stimuli used in priming studies may alter the processing level at which priming occurs. We also review recent behavioral, neuropsychological, and neurophysiological evidence that indicates that the priming patterns are reflected in activity modulations at multiple sites along the visual pathways. We furthermore suggest that studies of priming in visual search may potentially shed important light on the nature of cortical visual representations. Our conclusion is that priming occurs at many different levels of the perceptual hierarchy, reflecting activity modulations ranging from lower to higher levels, depending on the stimulus, task, and context—in fact, the neural loci that are involved in the analysis of the stimuli for which priming effects are seen.     

Cognitive neuroscience of episodic memory encoding

This paper presents a cognitive neuroscientific perspective on how human episodic memories are formed. Convergent evidence from multiple brain imaging studies using positron emission tomography (PET) and functional magnetic resonance imaging (fMRI) suggests a role for frontal cortex in episodic memory encoding. Activity levels within frontal cortex can predict episodic memory encoding across a wide range of behavioral manipulations known to influence memory performance, such as those present during levels of processing and divided attention manipulations. Activity levels within specific frontal and medial temporal regions can even predict, on an item by item basis, whether an episodic memory is likely to form. Furthermore, separate frontal regions appear to participate in supplying code-specific information, including distinct regions which process semantic attributes of verbal information as well as right-lateralized regions which process nonverbal information. We hypothesize that activity within these multiple frontal regions provides a functional influence (input) to medical temporal regions that bind the information together into a lasting episodic memory trace.     

Neurophysiology of successful encoding and retrieval of source memory

Event-related potentials were recorded during encoding, to identify whether brain activity predicts subsequent retrieval of spatial source information, and during retrieval, to investigate the neural correlates of successful and unsuccessful spatial context recollection. The amplitude registered during encoding for study items that were later recognized and assigned a correct source judgment was more positive than the amplitude for recognized items given incorrect source judgments; this difference started 480 msec poststimulus, predominantly at central and anterior sites. During retrieval, the waveform was more positive from 250 to 1,600 msec poststimulus when the brain had retrieved episodic information successfully than when it had failed. These findings indicate that brain electrical activity recorded during the first presentation of an item within its context predicts the subsequent retrieval of the specific spatial context. During retrieval, brain activity differed quantitatively at anterior sites and qualitatively at posterior sites according to the accuracy of source memory.     

Attention during memory retrieval enhances future remembering

Memory retrieval is a powerful learning event that influences whether an experience will be remembered in the future. Although retrieval can succeed in the presence of distraction, dividing attention during retrieval may reduce the power of remembering as an encoding event. In the present experiments, participants studied pictures of objects under full attention and then engaged in item recognition and source memory retrieval under full or divided attention. Two days later, a second recognition and source recollection test assessed the impact of attention during initial retrieval on long-term retention. On this latter test, performance was superior for items that had been tested initially under full versus divided attention. More importantly, even when items were correctly recognized on the first test, divided attention reduced the likelihood of subsequent recognition on the second test. The same held true for source recollection. Additionally, foils presented during the first test were also less likely to be later recognized if they had been encountered initially under divided attention. These findings demonstrate that attentive retrieval is critical for learning through remembering.     

Short-Term Visual Recognition Memory in Pigeons

Recognition memory for lists of items was investigated in pigeons using a YES-NO recognition technique. Experiment I showed that increasing the exposure duration of the first item of a two-item list improved recognition for that item without impairing recognition of the second item. Experiment II showed that decreasing the inter-trial interval had no effect on correct YES responses but significantly increased the number of false YES responses. Experiment III showed that recognition for the last two items of a three-item list was no poorer than that for lists of only two items. Experiment IV showed that increasing the delay between presentation and test of a two-item list (from 0·25-1 s) had a more disruptive effect on recognition for the second than for the first item. The data from these four experiments support a model proposed by Roberts and Grant, according to which memory traces are independent, and decay as a negatively accelerated function of time. Experiments V, VI, and VII investigated recognition for lists of three, four, and five items, and found no evidence for a primacy effect, performance being a linear function of time since sample offset.     

The effects of unitization on familiarity-based source memory: testing a behavioral prediction derived from neuroimaging data

Performance on tests of source memory is typically based on recollection of contextual information associated with an item. However, recent neuroimaging results have suggested that the perirhinal cortex, a region thought to support familiarity-based item recognition, may support source attributions if source information is encoded as a feature of the relevant item (i.e., "unitized"). The authors hypothesized that familiarity may contribute to source memory performance if item and source information are unitized during encoding, whereas performance may rely more heavily on recollection if source information is encoded as an arbitrary contextual association. Three source recognition experiments examining receiver operating characteristics and response deadline performance indicated that familiarity makes a greater contribution to source memory if source and item information are unitized during encoding. These findings suggest that familiarity can contribute to source recognition and that its contribution depends critically on the way item and source information are initially processed.     

Serial recall of colors: Two models of memory for serial order applied to continuous visual stimuli

This study investigated the effects of serial position and temporal distinctiveness on serial recall of simple visual stimuli. Participants observed lists of five colors presented at varying, unpredictably ordered interitem intervals, and their task was to reproduce the colors in their order of presentation by selecting colors on a continuous-response scale. To control for the possibility of verbal labeling, articulatory suppression was required in one of two experimental sessions. The predictions were derived through simulation from two computational models of serial recall: SIMPLE represents the class of temporal-distinctiveness models, whereas SOB-CS represents event-based models. According to temporal-distinctiveness models, items that are temporally isolated within a list are recalled more accurately than items that are temporally crowded. In contrast, event-based models assume that the time intervals between items do not affect recall performance per se, although free time following an item can improve memory for that item because of extended time for the encoding. The experimental and the simulated data were fit to an interference measurement model to measure the tendency to confuse items with other items nearby on the list—the locality constraint—in people as well as in the models. The continuous-reproduction performance showed a pronounced primacy effect with no recency, as well as some evidence for transpositions obeying the locality constraint. Though not entirely conclusive, this evidence favors event-based models over a role for temporal distinctiveness. There was also a strong detrimental effect of articulatory suppression, suggesting that verbal codes can be used to support serial-order memory of simple visual stimuli.     

An experimental analysis of memory processing

Rhesus monkeys were trained and tested in visual and auditory list-memory tasks with sequences of four travel pictures or four natural/environmental sounds followed by single test items. Acquisitions of the visual list-memory task are presented. Visual recency (last item) memory diminished with retention delay, and primacy (first item) memory strengthened. Capuchin monkeys, pigeons, and humans showed similar visual-memory changes. Rhesus learned an auditory memory task and showed octave generalization for some lists of notes--tonal, but not atonal, musical passages. In contrast with visual list memory, auditory primacy memory diminished with delay and auditory recency memory strengthened. Manipulations of interitem intervals, list length, and item presentation frequency revealed proactive and retroactive inhibition among items of individual auditory lists. Repeating visual items from prior lists produced interference (on nonmatching tests) revealing how far back memory extended. The possibility of using the interference function to separate familiarity vs. recollective memory processing is discussed.     

瞳孔变化在记忆加工中的生物标记作用

瞳孔在不同记忆任务的同一加工阶段内具有变化模式的一致性, 瞳孔的缩放变化反映了信息加工过程中神经系统的激活状态。在刺激呈现前的预编码阶段, 瞳孔随个体注意集中水平的提高而收缩。在信息编码阶段, 瞳孔扩张程度作为相继记忆效应的生物标记, 可预测信息未来提取阶段的回忆成绩。在信息提取阶段, 采用自由回忆任务时, 瞳孔随记忆负荷的释放而迅速收缩, 但每个项目的提取均会引发小幅度瞳孔扩张; 而在再认提取任务中, 呈现旧刺激比呈现新刺激时瞳孔扩张更明显, 产生瞳孔新旧效应。瞳孔是记忆加工中敏感而稳定的生物标记, 而瞳孔测量法是探究记忆加工生理机制的有效手段。未来可以从瞳孔追踪术的角度, 采用多种研究手段进一步探究记忆的生理机制。     

Selective attention meets spontaneous recognition memory: Evidence for effects at retrieval

Previous research on the effects of Divided Attention on recognition memory have shown consistent impairments during encoding but more variable effects at retrieval. The present study explored whether effects of Selective Attention at retrieval and subsequent testing were parallel to those of Divided Attention. Participants studied a list of pictures and then had a recognition memory test that included both full attention and selective attention (the to be responded to object was overlaid atop a blue outlined object) trials. All participants then completed a second recognition memory test. The results of 2 experiments suggest that subsequent tests consistently show impacts of the status of the ignored stimulus, and that having an initial test changes performance on a later test. The results are discussed in relation to effect of attention on memory more generally as well as spontaneous recognition memory research.     

Auditory list memory and interference processes in monkeys.

Memory of 2 rhesus monkeys (Macaca mulatta) was tested in a serial probe recognition task with lists of 4 natural or environmental sounds, different retention intervals, and different manipulations of interference. At short retention intervals, increasing the separation of list items reduced the primacy effect and produced a recency effect. Similar results were shown by increasing interference across lists through item repetitions or making the first 2 list items high-interference items. These results indicated that decreasing first-item performance reduced proactive interference on memory of the last list items. At long (20 s) retention intervals, making the last list items of high interference reduced the recency effect, reduced retroactive interference, and produced a primacy effect. Taken together, interference plays a role in determining the primacy and recency effects of the serial-position function.     

Developmental changes in memory encoding: insights from event‐related potentials

The aim of the present study was to investigate developmental changes in encoding processes between 6‐year‐old children and adults using event‐related potentials (ERPs). Although episodic memory (‘EM’) effects have been reported in both children and adults at retrieval and subsequent memory effects have been established in adults, no previous ERP studies have examined subsequent memory effects in children. This represents a critical gap in the literature because encoding processes, and changes in neural correlates supporting encoding, partially account for age‐related improvements in children's memory performance. Results revealed that subsequent memory effects differed between children and adults temporally, directionally, and topographically. These findings lend support to the hypothesis that encoding processes and their neural correlates are an important source of change in memory development. A video abstract of this article can be viewed at http://youtu.be/sH83_qVimgc .