DOUBLE DISSOCIATION BETWEEN MEMORY SYSTEMS UNDERLYING EXPLICIT AND IMPLICIT MEMORY IN THE HUMAN BRAIN

Abstract— Amnesic patients have impaired explicit memory that is evident in poor recall and recognition of words, yet can have intact implicit memory for words as measured by repetition priming, the enhanced efficiency for reprocessing those word's. The dissociation between explicit and implicit memory for words is a fundamental characteristic of normal cognition that could reflect two different functional architectures of the human brain two separate processing systems or two levels of operation of a single system with implicit memory less demanding of that system. We present a patient who has a lesion in the right occipital lobe and who showed intact explicit and impaired implicit memory for words. The deficit was specific to visual priming. The double dissociation between explicit and implicit visual memory for words indicates that separate processing systems mediate these two forms of memory, and that a memory system in right occipital cortex mediates implicit visual memory for words.     

Form-Specific Visual Priming in the Left and Right Hemispheres

Word fragment completion performance was examined for items that were presented in the same or different letter case at study and test. During the study phase words and nonwords were presented at central fixation, then during the test phase a divided visual field technique was used in which word fragments were presented briefly to the right hemisphere (left visual field) or the left hemisphere (right visual field). Previous research using the word stem completion task indicated that only the right hemisphere was sensitive to case changes in words from study to test. In contrast, the current results indicate that in the fragment completion task the priming effects for the test items presented to either hemisphere were greater when the fragments were in the same compared to different letter case at study and test. These results indicate that both hemispheres are capable of supporting form-specific visual implicit memory.     

Hemispheric asymmetry in a perceptual priming task: evidence from patients with unilateral brain damage.

The present study examined the relationship between unilateral brain damage and the effect of nonverbal perceptual priming using a picture-fragment completion task. Subjects consisted of 11 left brain-damaged patients, 8 right brain-damaged patients, and 10 healthy normal controls. The mean score of normal controls was significantly higher than those of left and right brain-damaged patients. Although there were significant effects of priming in all the groups, a significant difference in amplitude of priming effects was found only between right brain-damaged patients and normal controls. The correlation between amount of fragmentation at which there was identification and priming effect was not significant. We conclude that the right hemisphere is more involved in perceptual priming of form than is the left hemisphere, and form-dependent processing in the perceptual priming task has an asymmetrical distribution in the right and left hemispheres.     

Right to left hemispheric shift in occipital electroencephalographic responses to repeated Kimura figures

Gordon and Carmon (1976) reported that repeated presentations of an initially novel stimulus were associated with a transfer of cerebral dominance over time (trials) from the right to the left hemisphere. To test the generalizability of these results the proportions of alpha rhythms over the left and right occipital and parietal lobes were measured following the presentation of recurring and nonrecurring complex visual patterns (the Kimura Figures) to the upper or lower, left or right peripheral visual fields. Analysis showed increased electrical activation (as inferred by attenuated proportions of alpha rhythms) of the left occipital lobe but decreased activation of the right occipital lobe. This shift occurred during repeated presentations of the same stimuli but not during single presentations of different novel stimuli. There was no significant shift in this activity over the parietal lobes. These results are consistent with the reports of other researchers who have found a shift of dominant neuroelectrical activity from the right to the left hemisphere as the novelty of a visuospatial stimulus decreases.     

Dissociating perceptual and conceptual implicit memory in multiple sclerosis patients

Previous studies indicate that Multiple Sclerosis (MS) patients exhibit deficits in tests of explicit memory such as free recall, but show normal priming on implicit tests of memory such as word stem completion. However, the memory performance of patients with different MS disease subtypes has not been fully examined. In the current study, memory was assessed in Primary Progressive (PPMS), Relapsing Remitting (RRMS), and Secondary Progressive (SPMS) MS subgroups. Explicit memory as well as perceptual and conceptual implicit memory were examined using free recall, word fragment completion, and exemplar generation tests, respectively. All three groups of MS patients exhibited free recall deficits and normal priming on the exemplar generation test. However, the PPMS group exhibited a deficit in word fragment completion priming, whereas the RRMS and SPMS groups exhibited normal levels of priming on this task. Lesion load was assessed using magnetic resonance imaging and was negatively correlated with explicit memory performance, but it did not account for the observed deficits in perceptual implicit memory. The results indicate that PPMS patients exhibit a pattern of memory impairment that is distinct from that of the RRMS and SPMS groups. Moreover, the results indicate that perceptual implicit memory can be neurologically dissociated from conceptual implicit memory.     

Lateralized repetition priming for familiar faces: Evidence for asymmetric interhemispheric cooperation

Repetition priming refers to facilitated recognition of stimuli that have been seen previously. Although a great deal of work has examined the properties of repetition priming for familiar faces, little has examined the neuroanatomical basis of the effect. Two experiments are presented in this paper that combine the repetition priming paradigm with a divided visual field methodology to examine lateralized recognition of familiar faces. In the first experiment participants were presented with prime faces unilaterally to each visual field and target faces foveally. A significant priming effect was found for prime faces presented to the right hemisphere, but not for prime faces presented to the left hemisphere. In Experiment 2, prime and target faces were presented unilaterally, either to the same visual field or to the opposite visual field (i.e., either within hemisphere or across hemispheres). A significant priming effect was found for the within right hemisphere condition, but not for the within left hemisphere condition, replicating the findings of the first experiment. Priming was also found in both of the across hemispheres conditions, suggesting that interhemispheric cooperation occurs to aid recognition. Taken in combination these experiments provide two main findings. First, an asymmetric repetition priming effect was found, possibly as a result of asymmetric levels of activation following recognition of a prime face, with greater priming occurring within the right hemisphere. Second, there is evidence for asymmetric interhemispheric cooperation with transfer of information from the right hemisphere to the left hemisphere to facilitate recognition.     

Lateralized repetition priming for familiar faces: Evidence for asymmetric interhemispheric cooperation

Repetition priming refers to facilitated recognition of stimuli that have been seen previously. Although a great deal of work has examined the properties of repetition priming for familiar faces, little has examined the neuroanatomical basis of the effect. Two experiments are presented in this paper that combine the repetition priming paradigm with a divided visual field methodology to examine lateralized recognition of familiar faces. In the first experiment participants were presented with prime faces unilaterally to each visual field and target faces foveally. A significant priming effect was found for prime faces presented to the right hemisphere, but not for prime faces presented to the left hemisphere. In Experiment 2, prime and target faces were presented unilaterally, either to the same visual field or to the opposite visual field (i.e., either within hemisphere or across hemispheres). A significant priming effect was found for the within right hemisphere condition, but not for the within left hemisphere condition, replicating the findings of the first experiment. Priming was also found in both of the across hemispheres conditions, suggesting that interhemispheric cooperation occurs to aid recognition. Taken in combination these experiments provide two main findings. First, an asymmetric repetition priming effect was found, possibly as a result of asymmetric levels of activation following recognition of a prime face, with greater priming occurring within the right hemisphere. Second, there is evidence for asymmetric interhemispheric cooperation with transfer of information from the right hemisphere to the left hemisphere to facilitate recognition.     

Form-specific visual priming in the right cerebral hemisphere.

Results of 4 experiments indicate that both within-modality and case-specific visual priming for words are greater when test stimuli are presented initially to the right cerebral hemisphere (RH). In contrast, neither within-modality nor case-specific explicit memory for words is greater when stimuli are presented initially to the RH. Priming is measured using word-stem completion, and explicit memory is measured using word-stem cued recall. In both cases, Ss first rate how much they like words, and then word stems are presented briefly to the RH (in the left visual field) or to the left hemisphere (in the right visual field). Results suggest that at least 2 separate systems encode the visual representations that produce priming. The system that is more effective in the RH is better at representing form-specific information, whereas another system that is not more effective in the RH does not distinguish among distinct instances of word forms.     

Aging affects both perceptual and lexical/semantic components of word stem priming: An event-related fMRI study

In this event-related fMRI study, brain activity patterns were compared in extensive groups of young (N=25) and older (N=38) adults, while they were performing a word stem completion priming task. Based on behavioral findings, we tested the hypothesis that aging affects only the lexical/semantic, but not the perceptual component of word stem priming. To this end, we distinguished between priming-related activity reductions in posterior regions involved in visual processing, and regions associated with lexical/semantic retrieval processes, i.e., left lateral temporal and left prefrontal regions. Both groups revealed significant priming-related response time reductions. However, in accordance with earlier findings, a larger priming effect was found in the group of young participants. In line with previous imaging studies, the groups showed common priming-related activity reductions in the anterior cingulate, and the left inferior prefrontal cortex extending into the anterior portion of the left superior temporal gyrus, and at lower thresholds also in the right occipital lobe. However, when directly comparing the groups, greater priming-related reductions were found for the young group in the left anterior superior temporal gyrus and the right posterior occipital lobe. These findings suggest that, converse to current psychological views, aging affects both perceptual and lexical/semantic components of repetition priming.     

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     

Semantic and associative priming in the cerebral hemispheres: some words do, some words don't ... sometimes, some places

This study investigated spreading activation for words presented to the left and right hemispheres using an automatic semantic priming paradigm. Three types of semantic relations were used: similar-only (Deer-Pony), associated-only (Bee-Honey), and similar + associated (Doctor-Nurse). Priming of lexical decisions was symmetrical over visual fields for all semantic relations when prime words were centrally presented. However, when primes and targets were lateralized to the same visual field, similar-only priming was greater in the LVF than in the RVF, no priming was obtained for associated-only words, and priming was equivalent over visual fields for similar + associated words. Similar results were found using a naming task. These findings suggest that it is important to lateralize both prime and target information to assess hemisphere-specific spreading activation processes. Further, while spreading activation occurs in either hemisphere for the most highly related words (those related by category membership and association), our findings suggest that automatic access to semantic category relatedness occurs primarily in the right cerebral hemisphere. These results imply a unique role for the right hemisphere in the processing of word meanings. We relate our results to our previous proposal (Burgess & Simpson, 1988a; Chiarello, 1988c) that there is rapid selection of one meaning and suppression of other candidates in the left hemisphere, while activation spreads more diffusely in the right hemisphere. We also outline a new proposal that activation spreads in a different manner for associated words than for words related by semantic similarity.     

Cross-modal priming and explicit memory in patients with verbal production deficits

Implicit memory is often thought to reflect an influence of past experience on perceptual processes, yet priming effects are found when the perceptual format of stimuli changes between study and test episodes. Such cross-modal priming effects have been hypothesized to depend upon stimulus recoding processes whereby a stimulus presented in one modality is converted to other perceptual formats. The present research examined recoding accounts of cross-modal priming by testing patients with verbal production deficits that presumably impair the conversion of visual words into auditory/phonological forms. The patients showed normal priming in a visual stem completion task following visual study (Experiment 1), but showed impairments following auditory study in both implicit (Experiment 2) and explicit (Experiment 3) stem completion. The results are consistent with the hypothesis that verbal production processes contribute to the recoding of visual stimuli and support cross-modal priming. The results also indicate that shared processes contribute to both explicit memory and cross-modal implicit memory.     

RIGHT-HEMISPHERE MEMORY SUPERIORITY:

Abstract— Six experiments explored hemispheric memory differences in a patient who had undergone complete corpus callosum resection. The right hemisphere was better able than the left to reject new events similar to originally presented materials of several types, including abstract visual forms, faces, and categorized lists of words. Although the left hemisphere is capable of mental manipulation, imagination, semantic priming, and complex language production, these functions are apparently linked to memory confusions—confusions less apparent in the more literal right hemisphere. Differences between the left and right hemispheres in memory for new schematically consistent or categorically related events may provide a source of information allowing people to distinguish between what they actually witnessed and what they only inferred.     

Right hemispheric specialization for mental imagery: A review of the evidence

An implicit assumption in the literature on functional hemispheric asymmetry is that the right hemisphere plays a special role in mental imagery. Using a definition of mental imagery as quasi-perceptual experience, we draw distinctions among visual imagery, visual recognition memory, and visuospatial abilities. We then review the research literature to evaluate the hypothesis that mental imagery is a specialized function of the right cerebral hemisphere, and find that it receives little unambiguous support. Case reports of loss of imagery are no more frequent with right than with left unilateral brain damage. Systematic studies of brain-injured patients provide some support for the hypothesis, but are also consistent with the alternative hypothesis of bilateral representation of imagery. Commissurotomized patients report dreaming and being able to form visual images. Behavioral and psychophysiological studies of non-brain-injured patients either fail to provide evidence in favor of the hypothesis or can be interpreted as compatible with the alternate hypothesis of bilateral hemispheric involvement in imagery. We conclude that there is, presently, insufficient empirical basis for considering imagery a right hemispheric function. We then discuss implications of this conclusion for future research.     

Optic aphasia and the right hemisphere: a replication and extension.

Optic aphasia is a rare, visual modality-specific naming disorder. We describe a patient who developed this disorder subsequent to a left occipital lobe infarction. Like another case that we described previously, this patient performed normally on a variety of tasks assessing the recognition of objects he could not name. Additionally, although he never read aloud a single word, his performance on lexical decision and word comprehension tasks was far better than chance. We suggest that his performance was mediated by a right hemisphere semantic system.     

The effect of associative strength on priming in the cerebral hemispheres

Recent studies have suggested that semantic memory is more diffusely organized in the right hemisphere of the brain and that words directed to this hemisphere are more likely to activate meanings distantly related to the input. It is argued that this model of language processes predicts that variations in the associative strength of word pairs should give rise to different patterns of priming in each hemisphere. Specifically, the right hemisphere should exhibit relatively more facilitation than the left in response to weaker associative relationships, whereas the left hemisphere should exhibit relatively more facilitation than the right in the context of stronger relationships. This study varied the strength of the semantic association between prime and target in a divided visual field priming procedure. The results were unequivocal in demonstrating similar associative strength functions in each hemisphere, under conditions that encouraged either automatic or controlled processing. Visual field differences in absolute RTs to words and in magnitude of facilitation effects support the claim that the data collected in this study are veridical with respect to priming processes in the hemispheres. It is suggested that current models of hemispheric language processes require further refinement.     

Remembering 1500 pictures: the right hemisphere remembers better than the left

We hypothesized that the right hemisphere would be superior to the left hemisphere in remembering having seen a specific picture before, given its superiority in perceptually encoding specific aspects of visual form. A large set of pictures (N=1500) of animals, human faces, artifacts, landscapes, and art paintings were shown for 2s in central vision, or tachistoscopically (for 100ms) in each half visual field, to normal participants who were then tested 1-6 days later for their recognition. Images that were presented initially to the right hemisphere were better recognized than those presented to the left hemisphere. These results, obtained with participants with intact brains, large number of stimuli, and long retention delays, are consistent with previously described hemispheric differences in the memory of split-brain patients.     

额叶参与对新异联系的启动效应——来自脑损伤病人的证据

对新异联系的启动效应是内隐记忆的一种形式,又称联想启动。采用知觉辨认任务,探讨额叶损伤与联想启动的关系。25例额叶损伤病人及18例匹配的正常被试参与实验。结果表明,额叶受损病人辨认旧词对与重组词对的正确率没有明显差别;而且,轻度记忆损伤的额叶病人再认成绩正常,但联想启动值低于对照组,内隐记忆和外显数、坚持性反应数和语词流畅性等有中度相关。这些结果提示,额叶不仅与情节记忆、工作记忆等外显记忆有关,也参与了以非相关词对为材料的联想启动。     

A positron emission tomography (PET) study of autobiographical memory retrieval

Memory for the experiences of one's life, autobiographical memory (AM), is one of the most human types of memory, yet comparatively little is known of its neurobiology. A positron emission tomography (PET) study of AM retrieval revealed that the left frontal cortex was significantly active during retrieval (compared to memory control tasks), together with activation in the inferior temporal and occipital lobes in the left hemisphere. We propose that this left frontal lobe activation reflects the operation of control processes that modulate the construction of AMs in posterior neocortical networks.     

保留的跨通道启动与老化：对于近期观点的总结

通道内重复启动的研究提示老年人内隐记忆未受损, 这不只体现在视觉通道上还包括其他感觉通道(例如触觉、听觉和嗅觉)。然而很少有研究考察启动任务是否具有通道特异性。在以年轻人为被试的研究中发现跨通道迁移(视觉到触觉和触觉到视觉)和通道内迁移(视觉到视觉, 触觉到触觉)具有相似性。一项最近的研究进一步探索老年人在跨通道启动任务上是否受损。结果显示视觉和触觉间的跨通道启动在年轻被试和老年被试上都是保留的且具有对称性。并且, 对于自然声响、图片的通道内和跨通道启动任务随着年老化发展仍旧保留。这些行为结果和其它最近神经科学结果显示跨通道启动发生于枕叶后纹状皮层区, 而这一区域在老年人中未损坏。这一领域未来的研究方向包括利用不同知觉通道间、利用熟悉的和新异的刺激并结合行为的和脑成像的方法, 通过设计完善的跨通道启动来研究正常老人与阿尔兹海默病人, 还包括将设计得完善的启动任务包括在用于改善老年人记忆功能的项目中。