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The overall tripartite attribute model of memory is shown in Fig. 4. Different forms of memory and its neurobiological underpinnings are represented in terms of the nature, structure, or content of information representation as a set of different attributes, including language, time, place, response, reward value (affect), and visual object as an example of sensory perception. For each attribute, information is processed in the event-based memory system through operations that involve pattern separation or ortho-gonalization of specific attribute information, short-term memory processing, encoding of specific pattern associations into long-term memory, and retrieval of stored information via flexibility and pattern completion. In addition, for each attribute, information is processed in the knowledge-based system through operations of long-term storage, selective attention, perceptual memory, and retrieval of pattern associations. Finally, for each attribute, information is processed in the rule-based memory system through the integration of information from the event-based and knowledge-based memory systems for the selection of strategies and rules for maintaining or

Language

Sensory-Perception Response Reward Value (Affect) e.g. Visual Object

Language

Sensory-Perception Response Reward Value (Affect) e.g. Visual Object

Attribute

Place

Figure 4 Representation of the neural substrates associated with the event-based, knowledge-based, and rule-based memory systems for the language, time, place, response, reward value (affect), and sensory-perception attributes.

Attribute

Place

I I Rule-based Memory System f I Knowledge-based Memory System Event-based Memory System

Figure 5 Representation of the spatial attribute neural circuit incorporating neural regions that mediate rule-based, knowledge-based, and event-based memory.

I I Rule-based Memory System f I Knowledge-based Memory System Event-based Memory System

Figure 5 Representation of the spatial attribute neural circuit incorporating neural regions that mediate rule-based, knowledge-based, and event-based memory.

manipulating information for subsequent action. The neural systems that subserve specific attributes within a system can operate independent of each other, even though there are also many possibilities for interactions among the attributes. Although the event-based and knowledge-based memory systems are supported by neural substrates and different operating characteristics, suggesting that the two systems can operate independent of each other, there are also important interactions between the two systems, especially during the consolidation of new information and retrieval of previously stored information. Finally, because it is assumed that the rule-based system is influenced by the integration of event-based and knowledge-based memory information, there should be important interactions between the event-based and knowledge-based memory systems and the rule-based memory system. Thus, for each attribute there is a neural circuit that encompasses all three memory systems in representing specific attribute information. Space only allows for the presentation of one neural circuit as an example. Figure 5 depicts the neural substrates and their interconnections associated with the spatial (place) attribute across all three memory systems. Note that the dorsal lateral thalamus, pre- and parasubiculum, hippocampus, and subiculum represent neural substrates that support the event-based memory system; the entorhinal cortex, parahippocam-

pal gyrus or postrhinal cortex, posterior parietal cortex, and retrosplenial cortex support the knowledge-based memory system; and the lateral prefrontal cortex or pre- and infralimbic cortex support the rule-based memory system. This circuit provides anatomical support for a possible independence in the operation of the hippocampus as part of the event-based memory system and posterior parietal cortex as part of the knowledge-based memory system in that spatial information that is processed via the dorsal lateral thalamus can activate both the hippocampus and the posterior parietal cortex in parallel. Also, information can reach the lateral prefrontal cortex or pre- and infralimbic cortex as part of the rule-based memory system via direct connections from the posterior parietal cortex as part ofthe knowledge-based memory system and hippocampus as part of the event-based memory system. Finally, spatial information can interact with other specific attributes via a series of direct connections, including an interaction with reward value attribute information via hippocampal-amygdala connections or lateral prefrontal cortex-orbital frontal cortex connections or an interaction with response attribute information via hippocampal-caudate or lateral prefrontal-premotor or supplementary motor connections. In general, the tripartite attribute memory model represents the most comprehensive memory model capable of integrating the extant knowledge concerning the neural system representation of memory.

See Also the Following Articles

INFORMATION PROCESSING • LANGUAGE AND LEXICAL PROCESSING • MEMORY, EXPLICIT AND IMPLICIT • MEMORY DISORDERS, ORGANIC • MEMORY, NEUROIMAGING • MEMORY, OVERVIEW • SEMANTIC MEMORY • SHORT-TERM MEMORY • WORKING MEMORY

Suggested Reading

Burgess, N., Jeffery, K. J., and O'Keefe, J. (Eds.) (1999). The Hippocampal and Parietal Foundations of Spatial Cognition. Oxford Univ. Press, New York.

Eichenbaum, H. (1999). The hippocampus and mechanisms of declarative memory. Behav. Brain Res. 103, 123-133. Kesner, R. P. (1998). Neurobiological views of memory. In Neurobiology of Learning and Memory. (J. L. Martinez and R. P. Kesner, Eds.), pp. 361-416. Academic Press, New York. LeDoux, J. E. (1995). Emotion: Clues from the brain. Annu. Rev.

Psychol. 46, 209-235. Martinez, J. L., and Kesner, R. P. (Eds.) (1998). Neurobiology of

Learning and Memory. Academic Press, San Diego. Roberts, A. C., Robbins, T. W., and Weiskrantz, L. (Eds.) (1998). The Prefrontal Cortex: Executive and Cognitive Functions. Oxford Univ. Press, New York. Schacter, D. L., and Buckner, R. L. (1998). Priming and the brain.

Neuron 20, 185-195. Squire, L. R. (1994). Declarative and nondeclarative memory: Multiple brain systems supporting learning and memory. In Memory Systems 1994 (D. L. Schacter and E. Tulving, Eds.), pp. 203-231. MIT Press, Cambridge, MA.

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