Encoding the future: successful processing of intentions engages predictive brain networks.

Evidence from cognitive, patient and neuroimaging research indicates that "remembering to remember" intentions, i.e., prospective memory (PM) retrieval, requires both general memory systems involving the medial temporal lobes and an executive system involving rostral PFC (BA 10). However, it is not known how prospective memories are initially formed. Using fMRI, we investigated whether brain activity during encoding of future intentions and present actions differentially predicted later memory for those same intentions (PM) and actions (retrospective memory). We identified two significant patterns of neural activity: a network linked to overall memory and another linked specifically to PM. While overall memory success was predicted by temporal lobe activations that included the hippocampus, PM success was also uniquely predicted by activations in additional regions, including left rostrolateral PFC and the right parahippocampal gyrus. This finding extends the role of these structures to the formation of individual intentions. It also provides the first evidence that PM encoding, like PM retrieval, is supported by both a common episodic memory network and an executive network specifically recruited by future-oriented processing.

Encoding specificity revisited: the role of semantics.

Three experiments examined the effects of semantic characteristics of word pairs on memory using the encoding specificity paradigm. The paradigm involved four phases: (a) an encoding phase to relate cues and targets, (b) a phase in which words were generated to new cues, (c) a phase for recognition of generated targets, and (d) a cued-recall phase using the original encoding cues. Encoding pairs were classified a priori as either semantically similar (e.g., alluring-PRETTY), semantically contrasting (e.g., drab-PRETTY), or semantically unrelated (e.g., sore-PRETTY). Generation pairs were classified a priori as either semantically similar (e.g., beautiful-PRETTY) or semantically contrasting (e.g., ugly-PRETTY). For recall, the results showed that both the semantic relations between the encoding cue and target and the reprovision of the encoding cue at retrieval were important factors. In the case of recognition, however, both the semantic congruence between the encoding and generation contexts and the amount of semantic elaboration provided by the encoding context were important factors.

Parallel effects of aging and time pressure on memory for source: evidence from the spacing effect.

Two experiments address the degree to which item memory-the ability to remember that a word has been presented-is dissociable from source memory, or the ability to remember the context of that word's presentation. Spacing (as opposed to massing) an item's two presentations leads young adults to endorse that item more often when they are instructed to recognize it and to reject it more often when they are instructed to exclude it. Old adults also enjoy beneficial effects of spacing when the item is to be recognized, but suffer detrimental effects of the spacing manipulation when the item is to be rejected: They falsely endorse the spaced to-be-rejected items more than the massed ones. This dissociation also obtains with young subjects under conditions of increased time pressure: Under normal decision conditions, the ability to endorse to-be-recognized items and to reject to-be-rejected items increases with spacing; under speeded conditions, the ability to reject the latter items decreases with spacing. The results support the notion that source memory is selectively impaired in the elderly and that it is difficult to access mnemonic information about source under time pressure.

An examination of the effects of stimulus type, encoding task, and functional connectivity on the role of right prefrontal cortex in recognition memory.

Right anterior prefrontal cortex and other brain areas are active during memory retrieval but the role of prefrontal cortex and how it interacts with these other regions to mediate memory function remain unclear. To explore these issues we used positron emission tomography to examine the effects of stimulus material and encoding task on brain activity during visual recognition, assessing both task-related changes and functional connectivity. Words and pictures of objects were encoded using perceptual and semantic strategies, resulting in better memory for semantically encoded items. There was no significant effect of prior encoding strategy on brain activity during recognition. Right anterior prefrontal cortex was equally active during recognition of both types of stimuli irrespective of initial encoding strategy. Regions whose activity was positively correlated with activity in right anterior prefrontal cortex included widespread areas of prefrontal and inferior temporal cortices bilaterally. Activity in this entire network of regions was negatively correlated with recognition accuracy of semantically encoded items. These results suggest that initial encoding task has little impact on the set of brain regions that is active during subsequent recognition. Right anterior prefrontal cortex appears to be involved in retrieval mode, reflected in its equivalent activity across conditions differing in both stimulus type and encoding task, and also in retrieval effort, shown by the negative correlation between its functional connectivity and individual differences in recognition accuracy.

Attention and successful episodic encoding: an event-related potential study.

Event-related potentials (ERPs) were used to delineate the cerebral processes occurring when information is encoded into episodic memory and to determine how these processes are affected by divided attention. ERPs were recorded during encoding under focused or divided attention, and were selectively averaged on the basis of their retrieval during later free recall and recognition tests (with remember-know judgments). Items retrieved with conscious recollection of the encoding episode (remembered, recalled) were distinguished at encoding from later missed items by an enhanced left fronto-temporal negative wave (N340), a negative posterior sustained potential and a positive frontal sustained potential. These effects occurred independently of the level of attention. Items later retrieved on the basis of familiarity (known) elicited a larger N340 than missed items, but did not demonstrate the increased sustained potentials. We suggest that item-specific conceptual processing (N340) is sufficient to produce familiarity-based recognition, but additional elaborative processing (sustained interaction of frontal and posterior regions) is necessary for conscious recollection. The effect of divided attention on these processes was related to the difficulty of the secondary task, with the more difficult task causing greater and earlier interference.

Influences on the efficiency of prospective memory in younger and older adults.

The influence of 4 factors on age-related declines in prospective memory (PM) was considered in 2 experiments. The results of the experiments reveal that age-related differences in PM were not moderated by the degree of match between the nature of processing required in the ongoing activity and the defining features of the PM cue. Age-related differences in the accuracy of PM responses were primarily attributable to an increase in the number of omission errors committed by older adults. Age-related differences in PM were somewhat independent of the ability to recall the PM cues and intentions following task performance and were mediated by the cognitive resources of processing speed, inhibitory control, and working memory.

Asymmetry between encoding and retrieval processes: evidence from divided attention and a calibration analysis.

Two experiments provide further information on the effects of divided attention (DA) on encoding and retrieval processes. The first experiment examined the effects of decision and motor difficulty of a concurrent reaction time task. A calibration analysis was used in the second experiment to test the hypothesis that shifting attentional emphasis away from encoding to the secondary task reduces the level of processing the to-be-remembered items receive. Overall, the results confirm and extend the conclusions of Craik, Govoni, Naveh-Benjamin, and Anderson (1996) and Naveh-Benjamin, Craik, Guez, and Dori (1998), by pointing to clear differences between encoding and retrieval processes: Encoding is affected by simultaneous task demands, especially those associated with "central" resources involved in conscious decision making, whereas retrieval is obligatory in that it is largely immune to the effects of simultaneous demands. The results of the calibration analysis suggest that one reason for the poorer memory performance as a result of DA at encoding is a qualitative shift to less deep, elaborative strategies.

The effect of divided attention on memory for items and their context.

Memory for context, in comparison to memory for items, is a more demanding task and requires more attentional resources. We examined differences between item and context memory using divided attention at encoding and retrieval. Participants were presented with word lists and were instructed to learn the items (i.e., words), the intrinsic context (i.e., the colour of the cards on which each word was presented), and the extrinsic context (i.e., the temporal order of the words). Among 72 young adults, in comparison to conditions of full attention, divided attention applied at encoding only or retrieval only resulted in equally lower performance on all memory tasks; in contrast, divided attention applied at both encoding and retrieval resulted in lower performance only on memory for temporal order. The findings support the idea that memory for temporal order requires greater attentional resources and strategic processing than memory for items.

The effects of divided attention on encoding- and retrieval-related brain activity: A PET study of younger and older adults.

Divided attention (DA) disrupts episodic encoding, but has little effect on episodic retrieval. Furthermore, normal aging is associated with episodic memory impairments, and when young adults are made to encode information under DA conditions, their memory performance is reduced and resembles that of old adults working under full attention (FA) conditions. Together, these results suggest a common neurocognitive mechanism by which aging and DA during encoding disrupt memory performance. In the current study, we used PET to investigate younger and older adults' brain activity during encoding and retrieval under FA and DA conditions. In FA conditions, the old adults showed reduced activity in prefrontal regions that younger adults activated preferentially during encoding or retrieval, as well as increased activity in prefrontal regions young adults did not activate. These results indicate that prefrontal functional specificity of episodic memory is reduced by aging. During encoding, DA reduced memory performance, and reduced brain activity in left-prefrontal and medial-temporal lobe regions for both age groups, indicating that DA during encoding interferes with encoding processes that lead to better memory performance. During retrieval, memory performance and retrieval-related brain activity were relatively immune to DA for both age groups, suggesting that DA during retrieval does not interfere with the brain systems necessary for successful retrieval. Finally, left inferior prefrontal activity was reduced similarly by aging and by DA during encoding, suggesting that the behavioral correspondence between these effects is the result of a reduced ability to engage in elaborate encoding operations.

The effects of divided attention on encoding and retrieval processes: the resiliency of retrieval processes.

We have recently cast doubt (Craik, Govoni, Naveh-Benjamin, & Anderson, 1996; Naveh-Benjamin, Craik, Guez, & Dori, 1998) on the view that encoding and retrieval processes in human memory are similar. Divided attention at encoding was shown to reduce memory performance significantly, whereas divided attention at retrieval affected memory performance only minimally. In this article we examined this asymmetry further by using more difficult retrieval tasks, which require substantial effort. In one experiment, subjects had to encode and retrieve lists of unfamiliar name-nouns combinations attached to people's photographs, and in the other, subjects had to encode words that were either strong or weak associates of the cues presented with them and then to retrieve those words with either intra- or extra-list cues. The results of both experiments showed that unlike division of attention at encoding, which reduces memory performance markedly, division of attention at retrieval has almost no effect on memory performance, but was accompanied by an increase in secondary-task cost. Such findings again illustrated the resiliency of retrieval processes to manipulations involving the withdrawal of attention. We contend that retrieval processes are obligatory or protected, but that they require attentional resources for their execution.

The effect of divided attention on encoding and retrieval in episodic memory revealed by positron emission tomography.

The effects of divided attention (DA) on episodic memory encoding and retrieval were investigated in 12 normal young subjects by positron emission tomography (PET). Cerebral blood flow was measured while subjects were concurrently performing a memory task (encoding and retrieval of visually presented word pairs) and an auditory tone-discrimination task. The PET data were analyzed using multivariate Partial Least Squares (PLS), and the results revealed three sets of neural correlates related to specific task contrasts. Brain activity, relatively greater under conditions of full attention (FA) than DA, was identified in the occipital-temporal, medial, and ventral-frontal areas, whereas areas showing relatively more activity under DA than FA were found in the cerebellum, temporo-parietal, left anterior-cingulate gyrus, and bilateral dorsolateral-prefrontal areas. Regions more active during encoding than during retrieval were located in the hippocampus, temporal and the prefrontal cortex of the left hemisphere, and regions more active during retrieval than during encoding included areas in the medial and right-prefrontal cortex, basal ganglia, thalamus, and cuneus. DA at encoding was associated with specific decreases in rCBF in the left-prefrontal areas, whereas DA at retrieval was associated with decreased rCBF in a relatively small region in the right-prefrontal cortex. These different patterns of activity are related to the behavioral results, which showed a substantial decrease in memory performance when the DA task was performed at encoding, but no change in memory levels when the DA task was performed at retrieval.

Divided attention during encoding and retrieval: differential control effects?

Previous studies have shown that divided attention (DA) during retrieval has little effect on recall of episodic memories, although DA during encoding has a large detrimental effect. One possible reason for this asymmetry is that stimulus presentation at encoding is under experimenter control, whereas retrieval operations and responses are under participant control. This experiment tested this possibility by presenting paired-associate word lists for learning and recall, either at a fixed 4-s rate or at a rate controlled by the participant. The results showed that the higher recall levels for DA at retrieval than for DA at encoding held under all combinations of experimenter and participant control. The implications of these results for a fuller understanding of encoding and retrieval processes are discussed.

Changes in memory processing with age.

Over the years, a large body of literature has shown that humans display losses in memory with age, but that not all types of memory are affected equally. Similarly, recent evidence from functional neuroimaging experiments has revealed that, depending on the task, older adults can display greater or lesser activity in task-relevant brain areas compared with younger adults. Recent behavioral and neurophysiological experiments are furthering our understanding of the effects of aging on cognition. It appears that some brain changes seen with age may be compensatory.

The effects of age on the neural correlates of episodic encoding.

Young and old adults underwent positron emission tomographic scans while encoding pictures of objects and words using three encoding strategies: deep processing (a semantic living/nonliving judgement), shallow processing (size judgement) and intentional learning. Picture memory exceeded word memory in both young and old groups, and there was an age-related decrement only in word recognition. During the encoding tasks three brain activity patterns were found that differentiated stimulus type and the different encoding strategies. The stimulus-specific pattern was characterized by greater activity in extrastriate and medial temporal cortices during picture encoding, and greater activity in left prefrontal and temporal cortices during encoding of words. The older adults showed this pattern to a significantly lesser degree. A pattern distinguishing deep processing from intentional learning of words and pictures was identified, characterized mainly by differences in prefrontal cortex, and this pattern also was of significantly lesser magnitude in the old group. A final pattern identified areas with increased activity during deep processing and intentional learning of pictures, including left prefrontal and bilateral medial temporal regions. There was no group difference in this pattern. These results indicate age-related dysfunction in several encoding networks, with sparing of one specifically involved in more elaborate encoding of pictures. These age-related changes appear to affect verbal memory more than picture memory.

Source memory and divided attention: reciprocal costs to primary and secondary tasks.

Source memory, in comparison with item memory, is more sensitive to frontal lesions and may require more strategic processing. Divided attention was used to restrict attentional resources and strategic processing on memory tasks. Participants encoded and retrieved items (i.e., words) and source (i.e., voice or spatial location) while concurrently performing a finger-tapping (FT) or visual reaction-time (VRT) task. Memory accuracy costs under divided attention were greater for retrieval of source than item and were greater with VRT than FT. Similarly, costs to the secondary task were greater when concurrently retrieving source as opposed to item and were greater for VRT than FT. Effects were stronger when spatial location was used as the source task. Findings support the idea that processing source information requires more attentional resources and effort than processing item information. Furthermore, concurrent performance of VRT produced greater interference with a task that was more dependent on intact frontal functioning and better simulated the performance of patients with frontal dysfunction.

Dissociation of attentional processes in patients with focal frontal and posterior lesions.

A location-based ('select-what, respond-where') priming task was used to examine three measures of selective attention (interference (INT), negative priming (NP), and inhibition of return (IOR)) as a function of focal brain pathology and the complexity of target selection. Control subjects showed different patterns of performance for the three attentional measures as a function of complexity, suggesting some independence among INT, NP, and IOR. Brain-damaged subjects showed significant response slowing, as well as a number of lesion-specific attentional abnormalities. Right frontal (including bifrontal) damage resulted in proportionally increased interference related to task complexity. Left posterior damage increased IOR in the most complex task, while left frontal damage reversed the control pattern of IOR as a function of complexity. Right hemisphere (right posterior and right frontal damage) pathology resulted in a virtual loss of negative priming at all levels of task complexity; left and bifrontal damage resulted in diminished NP only related to increases in the complexity of selection. INT, NP, and IOR are mediated by different brain regions and their expression can be modulated by the complexity of the selection task.

Age-related decline in prospective memory: the roles of cue accessibility and cue sensitivity.

In two experiments the authors evaluated the hypothesis that age-related decline in prospective memory reflects momentary lapses of intention (MLIs), and explored two factors, cue sensitivity and accessibility, that may contribute to MLIs. MLIs were reliably greater than zero in Experiment 1, indicating that performance fluctuated over the course of the task. Analysis of the response latency data (RL) revealed that older adults demonstrated elevated RL for missed prospective cues and were much slower to respond correctly to prospective cues than younger adults. These findings indicate preserved cue sensitivity in later adulthood and an age-related decline in cue accessibility. Experiment 2 demonstrated that cue sensitivity did not result from an orienting response to the perceptual novelty associated with the prospective cues.

Aging and judgments of duration: effects of task complexity and method of estimation.

The effects of aging on judgments of short temporal durations were explored using the prospective paradigm and the methods of verbal estimation and production. Younger and older adults performed a perceptual judgment task at five levels of complexity for periods of 30, 60, and 120 sec. Participants either continued to perform the task for a specified interval (production) or were stopped and then verbally estimated the interval. Older adults gave shorter verbal estimates and longer productions than did younger adults. The methods of verbal estimation and production yielded approximately equal duration judgment ratios once range effects were taken into account. Task complexity had little effect. The major conclusion is that duration judgment ratios decrease from younger to older adults when the intervals are filled with a mental task.

The attentional demands of encoding and retrieval in younger and older adults: 1. Evidence from divided attention costs.

Four studies examined the effects of divided attention in younger and older adults. Attention was divided at encoding or retrieval in free recall (Experiment 1), cued recall (Experiments 2 and 3), and recognition (Experiment 4). Dividing attention at encoding disrupted memory performance equally for the two age groups; by contrast, for both age groups, dividing attention at retrieval had little or no effect on memory performance. Secondary task reaction times (RTs) were slowed to a greater extent for the older adults than for the younger adults, especially at retrieval. Age-related differences in RT costs at retrieval were largest in free recall, smaller in cued recall, and smallest in recognition. These results provide evidence for an age-related increase in the attentional demands of encoding and retrieval.