Large,colorful, or noisy? Attribute- and modality-specific activations during retrieval of perceptual attribute knowledge

Position emission tomography was used to investigate whether retrieval of perceptual knowledge from long-term memory activates unique cortical regions associated with the modality and/or attribute type retrieved. Knowledge about the typical color, size, and sound of common objects and animals was probed, in response to written words naming the objects. Relative to a nonsemantic control task, all the attribute judgments activated similar left temporal and frontal regions. Visual (color, size) knowledge selectively activated the right posterior inferior temporal (PIT) cortex, whereas sound judgments elicited selective activation in the left posterior superior temporal gyrus and the adjacent parietal cortex. All of the attribute judgments activated a left PIT region, but color retrieval generated more activation in this area. Size judgments activated the right medial parietal cortex. These results indicate that the retrieval of perceptual semantic information activates not only a general semantic network, but also cortical areas specialized for the modality and attribute type of the knowledge retrieved.     

Convergent cortical representation of semantic processing in bilinguals

This study examined whether semantic processes in two languages (English and Spanish) are mediated by a common neural system in fluent bilinguals who acquired their second language years after acquiring their first language. Functional magnetic resonance imaging was performed while bilingual participants made semantic and nonsemantic decisions about words in Spanish and English. There was greater activation for semantic relative to nonsemantic decisions in left and right frontal regions, with greater left frontal activation. The locations of activations were similar for both languages, and no differences were found when semantic decisions for English and Spanish words were compared directly. These results demonstrate a shared frontal lobe system for semantic analysis of the languages and are consistent with cognitive research on bilingualism indicating that the two languages of a bilingual person access a common semantic system.     

Adult age differences in visual word identification: functional neuroanatomy by positron emission tomography

Adult age differences in the neural systems mediating semantic (context-independent) memory were investigated using positron emission tomography (PET). Younger (20-29 years) and older (62-70 years) participants performed lexical decision (word/nonword discrimination) and nonsemantic (simple visual search) baseline tasks during PET scanning. Within the lexical decision task, display duration and presentation rate were varied across scans. The behavioral data suggested that although an age-related slowing was evident in visual feature and response processing, the retrieval of semantic/lexical information was similar for younger and older adults. For both age groups, lexical-related activation occurred in inferior prefrontal and occipitotemporal regions of the left hemisphere. Differential activation, as a function of age group, was observed in the left occipitotemporal pathway as a result of older adults' maintaining higher levels of neural activity in striate cortex (during visual search) and in inferior temporal cortex (during lexical decision). The prefrontal activation was similar for the two age groups. Thus, although this form of semantic memory retrieval does not undergo significant age-related decline, an age-related change in the associated pattern of neural activation is evident. These findings differ from previous neuroimaging studies of episodic (context-dependent) memory retrieval, which have suggested that age-related compensatory mechanisms are expressed primarily by greater activation of prefrontal regions for older adults than for younger adults.     

Attention and the acquisition of new knowledge: their effects on older adults' associative memory deficit

Older adults experience a selective associative memory deficit by demonstrating intact item memory relative to impaired associative memory when compared with younger adults. Age-related deficits in associative memory have been suggested to arise from declines in attentional resources, and the role of attention during encoding and retrieval in associative memory for words and their spatial locations was investigated in the current experiment. Additionally, the tendency of younger and older adults to use knowledge acquired during encoding to improve their associative memory judgments through a strategic associative memory process was also investigated. Younger and older adults studied a list of words with each word belonging to one of four categories, which followed one of four mathematical probability structures for their presentation. Older adults exhibited intact item memory and impaired associative memory relative to full attention younger adults. In addition, both older and younger adults demonstrated an ability to engage in strategic associative memory, by learning and later using the probability structure introduced at study to guide their associative memory judgments. In contrast, dividing the attention of younger adults during encoding impaired item memory, associative memory and strategic associative memory, whereas dividing attention at retrieval did not result in similar deficits. These data add to a growing body of literature demonstrating older adults to exhibit a selective associative memory deficit that is not simulated by dividing the attention of younger adults at encoding or retrieval. Furthermore, younger and older adults maintain the ability to use new knowledge to guide their associative judgments.     

Effects of level of retrieval success on recall-related frontal and medial temporal lobe activations

Brain dedicated single photon emission computed tomography (SPECT) was used to compare the neuroactivation produced by the cued recall of response words in a set of studied word pairs with that produced by the cued retrieval of words semantically related to unstudied stimulus words. Six of the 12 subjects scanned were extensively trained so as to have good memory of the studied pairs and the remaining six were minimally trained so as to have poor memory. When comparing episodic with semantic retrieval, the well-trained subjects showed significant left medial temporal lobe activation, which was also significantly greater than that shown by the poorly trained subjects, who failed to show significant medial temporal lobe activation. In contrast, the poorly trained subjects showed significant bilateral frontal lobe activation, which was significantly greater than that shown by the well-trained subjects who failed to show significant frontal lobe activation. The frontal activations occurred mainly in the dorsolateral region, but extended into the ventrolateral and, to a lesser extent, the frontal polar regions. It is argued that whereas the medial temporal lobe activation increased as the proportion of response words successfully recalled increased, the bilateral frontal lobe activation increased in proportion to retrieval effort, which was greater when learning had been less good.     

IMPLICIT MEMORY IN AMNESIC PATIENTS:

Abstract— Previous observations of spared priming in amnesic patients have been based almost entirely on data from visual implicit memory tests Our research examined perceptual priming.in amnesic patients and control subjects on an auditory identification task in which previously spoken words and new words were presented in white noise We manipulated type of encoding task (semantic vs nonsemantic) and speaker's voice at study and test (same vs different) Priming was little affected.by either manipulation, and amnesic patients exhibited normal priming in all experimental conditions On an explicit test of recognition memory, by contrast, amnesic patients exhibited.severely impaired performance following the semantic study task, all subjects showed poor explicit memory following the nonsemantic study task Results are consistent with the idea that auditory priming depends largely on a presemantic auditory perceptual representation system     

Value-based modulation of memory encoding involves strategic engagement of fronto-temporal semantic processing regions

A number of prior fMRI studies have focused on the ways in which the midbrain dopaminergic reward system coactivates with hippocampus to potentiate memory for valuable items. However, another means by which people could selectively remember more valuable to-be-remembered items is to be selective in their use of effective but effortful encoding strategies. To broadly examine the neural mechanisms of value on subsequent memory, we used fMRI to assess how differences in brain activity at encoding as a function of value relate to subsequent free recall for words. Each word was preceded by an arbitrarily assigned point value, and participants went through multiple study–test cycles with feedback on their point total at the end of each list, allowing for sculpting of cognitive strategies. We examined the correlation between value-related modulation of brain activity and participants’ selectivity index, which measures how close participants were to their optimal point total, given the number of items recalled. Greater selectivity scores were associated with greater differences in the activation of semantic processing regions, including left inferior frontal gyrus and left posterior lateral temporal cortex, during the encoding of high-value words relative to low-value words. Although we also observed value-related modulation within midbrain and ventral striatal reward regions, our fronto-temporal findings suggest that strategic engagement of deep semantic processing may be an important mechanism for selectively encoding valuable items.     

Judgments of origin and generation effects: comparisons between young and elderly adults

In 2 experiments, young and elderly adults were required to both read words, and generate words by completing word fragments. Subjects were then required to recognize those words that had been presented earlier; for those words that they recognized they judged whether the items had initially been presented in read or generate form. Generation effects (better memory for words that were generated as compared with words that were read) of similar magnitude were observed for both young and older adults. The older adults were consistently less accurate than the younger adults in their judgments of origin. In addition, the young adults exhibited a bias to respond "read" for these judgments. In contrast, the older adults either exhibited a neutral response bias or were biased to respond "generate." Age-related differences in the encoding or retrieval of information about cognitive operations do not provide a good account of the results. Alternative accounts are described.     

Aging and semantic cueing during learning and retention of verbal episodic information

The purpose of this study was to determine the effectiveness of semantic cues provided at encoding and during retention for older adults' memory. For the California Verbal Learning Test-II, participants received semantic or nonsemantic cues that were varied across groups at encoding and during the retention interval. Provision of a semantic cue at encoding led to greater semantic clustering at learning, but not increased recall performance. Providing a semantic cue during the retention interval led to better delayed free recall and greater semantic clustering. No group differences in recall or semantic clustering were found at delayed cued recall. The current findings suggest that semantic cues can be beneficial for recalling unstructured information when administered during the retention interval.     

Integrative and semantic relations equally alleviate age-related associative memory deficits

Two experiments compared effects of integrative and semantic relations between pairs of words on lexical and memory processes in old age. Integrative relations occur when two dissimilar and unassociated words are linked together to form a coherent phrase (e.g., horse-doctor). In Experiment 1, older adults completed a lexical-decision task where prime and target words were related either integratively or semantically. The two types of relation both facilitated responses compared to a baseline condition, demonstrating that priming can occur in older adults with minimal preexisting associations between primes and targets. In Experiment 2, young and older adults completed a cued recall task with integrative, semantic, and unrelated word pairs. Both integrative and semantic pairs showed significantly smaller age differences in associative memory compared to unrelated pairs. Integrative relations facilitated older adults' memory to a similar extent as semantic relations despite having few preexisting associations in memory. Integratability of stimuli is therefore a new factor that reduces associative deficits in older adults, most likely by supporting encoding and retrieval mechanisms.     

Individual differences in skilled adult readers reveal dissociable patterns of neural activity associated with component processes of reading

We used fMRI to examine patterns of brain activity associated with component processes of visual word recognition and their relationships to individual differences in reading skill. We manipulated both the judgments adults made on written stimuli and the characteristics of the stimuli. Phonological processing led to activation in left inferior frontal and temporal regions whereas semantic processing was associated with bilateral middle frontal activation. Individual differences in reading subskills were reflected in differences in the degree to which cortical regions were engaged during reading. Variation in sight word reading efficiency was associated with degree of activation in visual cortex. Increased phonological decoding skill was associated with greater activation in left temporo-parietal cortex. Greater reading comprehension ability was associated with decreased activation in anterior cingulate and temporal regions. Notably, associations between reading ability and neural activation indicate that brain/behavior relationships among skilled readers differ from patterns associated with dyslexia and reading development.     

Semantic versus phonological false recognition in aging and Alzheimer's disease

Patients with Alzheimer's disease (AD) have been found to exhibit lower levels of false recognition of semantic associates compared with healthy older adults. Because these patients may show impaired performance of episodic and semantic memory tasks, this finding could be explained by deficits in episodic memory, semantic memory, or both. The authors adapted a paradigm for comparison of semantic versus phonological false recognition. They found that: (a) patients with AD exhibited lower levels of corrected false recognition of semantic, phonological, and hybrid (mixed semantic and phonological) lists than older adults, and (b) patients with AD showed very similar levels of false recognition for all list types. These results suggest that only episodic memory deficits are necessary to explain the lower level of false recognition of semantic associates observed in patients with AD when compared to older adults. Additionally, (c) older adults showed greater levels of semantic, phonological, and hybrid false recognition than younger adults, extending previous false recognition research of semantically related words and categorized colored photographs to phonologically related words.     

Frontotemporal neural systems supporting semantic processing in Alzheimer’s disease

We hypothesized that semantic memory for object concepts involves both representations of visual feature knowledge in modality-specific association cortex and heteromodal regions that are important for integrating and organizing this semantic knowledge so that it can be used in a flexible, contextually appropriate manner. We examined this hypothesis in an fMRI study of mild Alzheimer’s disease (AD). Participants were presented with pairs of printed words and asked whether the words matched on a given visual–perceptual feature (e.g., guitar, violin: SHAPE). The stimuli probed natural kinds and manufactured objects, and the judgments involved shape or color. We found activation of bilateral ventral temporal cortex and left dorsolateral prefrontal cortex during semantic judgments, with AD patients showing less activation of these regions than healthy seniors. Moreover, AD patients showed less ventral temporal activation than did healthy seniors for manufactured objects, but not for natural kinds. We also used diffusion-weighted MRI of white matter to examine fractional anisotropy (FA). Patients with AD showed significantly reduced FA in the superior longitudinal fasciculus and inferior frontal-occipital fasciculus, which carry projections linking temporal and frontal regions of this semantic network. Our results are consistent with the hypothesis that semantic memory is supported in part by a large-scale neural network involving modality-specific association cortex, heteromodal association cortex, and projections between these regions. The semantic deficit in AD thus arises from gray matter disease that affects the representation of feature knowledge and processing its content, as well as white matter disease that interrupts the integrated functioning of this large-scale network.     

Aging and Semantic Cueing during Learning and Retention of Verbal Episodic Information

ABSTRACT

The purpose of this study was to determine the effectiveness of semantic cues provided at encoding and during retention for older adults' memory. For the California Verbal Learning Test-II, participants received semantic or nonsemantic cues that were varied across groups at encoding and during the retention interval. Provision of a semantic cue at encoding led to greater semantic clustering at learning, but not increased recall performance. Providing a semantic cue during the retention interval led to better delayed free recall and greater semantic clustering. No group differences in recall or semantic clustering were found at delayed cued recall. The current findings suggest that semantic cues can be beneficial for recalling unstructured information when administered during the retention interval.     

Orienting task effects on memory for presentation modality in children, young adults, and older adults.

Children (7 to 10 years), young adults (17 to 24 years), and older adults (55 to 77 years) were asked to learn three lists of words that were of mixed modality (half the words were visual, and half the words were auditory). With one list the subjects were asked a semantic orienting question; with another, a nonsemantic orienting question; and with a third, no orienting question. Half the subjects in each age group were also asked to remember the presentation modality of each word. Older adults remembered less information about modality than children and young adults did, and the variation in the type of orienting question--or the lack of one--affected modality identification. However, there was no Orienting Task x Age interaction for modality identification. The results of this study suggest that encoding modality information does not take place automatically--in any age group--but that explanations focusing on encoding strategies and effort are not likely to account for older adults' difficulties in remembering presentation modality.     

Episodic encoding and recognition of pictures and words: role of the human medial temporal lobes

In the present PET study, we examined brain activity related to processing of pictures and printed words in episodic memory. Our goal was to determine how the perceptual format of objects (verbal versus pictorial) is reflected in the neural organization of episodic memory for common objects. We investigated this issue in relation to encoding and recognition with a particular focus on medial temporal-lobe (MTL) structures. At encoding, participants saw pictures of objects or their written names and were asked to make semantic judgments. At recognition, participants made yes-no recognition judgments in four different conditions. In two conditions, target items were pictures of objects; these objects had originally been encoded either in picture or in word format. In two other conditions, target items were words; they also denoted objects originally encoded either as pictures or as words. Our data show that right MTL structures are differentially involved in picture processing during encoding and recognition. A posterior MTL region showed higher activation in response to the presentation of pictures than of words across all conditions. During encoding, this region may be involved in setting up a representation of the perceptual information that comprises the picture. At recognition, it may play a role in guiding retrieval processes based on the perceptual input, i.e. the retrieval cue. Another more anterior right MTL region was found to be differentially involved in recognition of objects that had been encoded as pictures, irrespective of whether the retrieval cue provided was pictorial or verbal in nature; this region may be involved in accessing stored pictorial representations. Our results suggest that left MTL structures contribute to picture processing only during encoding. Some regions in the left MTL showed an involvement in semantic encoding that was picture specific; others showed a task-specific involvement across pictures and words. Together, our results provide evidence that the involvement of some but not all MTL regions in episodic encoding and recognition is format specific.     

Lateral and medial prefrontal contributions to emotion generation by semantic elaboration during episodic encoding

Memories for emotion-laden stimuli are remembered more accurately than those for neutral stimuli. Although this enhancement reflects stimulus-driven modulation of memory by emotions, functional neuroimaging evidence of the interacting mechanisms between emotions generated by intentional processes, such as semantic elaboration, and memory is scarce. The present fMRI study investigated how encoding-related activation is modulated by emotions generated during the process of semantic elaboration. During encoding with fMRI, healthy young adults viewed neutral (target) pictures either passively or with semantic elaboration. In semantic elaboration, participants imagined background stories related to the pictures. Encoding trials with semantic elaboration were subdivided into conditions in which participants imagined negative, positive, or neutral stories. One week later, memories for target pictures were tested. In behavioral results, memories for target pictures were significantly enhanced by semantic elaboration, compared to passive viewing, and the memory enhancement was more remarkable when negative or positive stories were imagined. fMRI results demonstrated that activations in the left inferior frontal gyrus and dorsal medial prefrontal cortex (dmPFC) were greater during the encoding of target pictures with semantic elaboration than those with passive viewing, and that these activations further increased during encoding with semantic elaboration of emotional stories than of neutral stories. Functional connectivity between the left inferior frontal gyrus and dmPFC/hippocampus during encoding significantly predicted retrieval accuracies of memories encoded with self-generated emotional stories. These findings suggest that networks including the left inferior frontal region, dmPFC, and hippocampus could contribute to the modulation of memories encoded with the emotion generation.     

FUNCTIONAL MAGNETIC RESONANCE IMAGING OF SEMANTIC MEMORY PROCESSES IN THE FRONTAL LOBES

Abstract— Frontal-lobe activation during semantic memory performance was examined using functional magnetic resonance imaging (fMRI), a noninvasive technique for localizing neural activity associated with cognitive function. Left inferior prefrontal cortex was more activated for semantic than for perceptual encoding of words, and for initial than for repeated semantic encoding of words. Decreased activation for semantic encoding of repeated words reflects repetition priming, that is, implicit retrieval of memory gained in the initial semantic encoding of a word. The left inferior prefrontal region may sub-serve semantic working memory processes that participate in semantic encoding and that have decreased demands when such encoding can be facilitated by recent semantic experience. These results demonstrate that fMRI can visualize changes in an individual's brain function associated with the encoding and retrieval of new memories.     

Breaking the waves: age differences in electrical brain activity when reading text with distractors

The authors tested whether older adults have greater difficulty than younger adults in ignoring task-irrelevant information during reading as a result of age-related decline in inhibitory processes. Participants were shown target sentences containing distractor words. They were instructed to read aloud each sentence and ignore distractors. The N400 event-related potential (ERP) was used to measure the extent of semantic processing of target and distracting information. It showed that younger adults semantically processed both target and distracting material, whereas online processing of target sentences in older adults was disrupted by the distractors. In older adults, memory for target information related to their susceptibility to distraction and inhibition efficiency. Implications for age-differences in inhibitory control, working memory, and resource capacity are discussed.     

Perceptual specificity and implicit auditory priming in older and younger adults.

Age differences in perceptual specificity for implicit auditory priming were examined in 3 experiments. All 3 experiments began with a study phase during which participants rated words based on perceptual (shallow encoding) or semantic (deep encoding) attributes. After the study phase, participants were asked to identify filtered versions of repeated and new words (implicit test) and then to make old/new recognition judgments (explicit test). In contrast to earlier findings (D. L. Schacter, B. Church, & D. M. Osowiecki, 1994), older and younger adults were equally sensitive to study-to-test changes in speaking rate (Experiment 1), fundamental frequency (Experiment 2), and voice (Experiment 3). Explicit memory, in contrast, was significantly poorer for older adults but was minimally affected by changes in surface features. Findings from the study are discussed with respect to their implications for establishing the mechanisms mediating perceptual specificity and for their importance in understanding age-related changes in implicit memory.