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- Daniel S. Ruchkin, Jordan Grafman, Katherine Cameron & Rita S. Berndt (2003). Working Memory Retention Systems: A State of Activated Long-Term Memory. Behavioral and Brain Sciences 26 (6):709-728.High temporal resolution event-related brain potential and electroencephalographic coherence studies of the neural substrate of short-term storage in working memory indicate that the sustained coactivation of both prefrontal cortex and the posterior cortical systems that participate in the initial perception and comprehension of the retained information are involved in its storage. These studies further show that short-term storage mechanisms involve an increase in neural synchrony between prefrontal cortex and posterior cortex and the enhanced activation of long-term memory representations of material held in short-term memory. This activation begins during the encoding/comprehension phase and evidently is prolonged into the retention phase by attentional drive from prefrontal cortex control systems. A parsimonious interpretation of these findings is that the long-term memory systems associated with the posterior cortical processors provide the necessary representational basis for working memory, with the property of short-term memory decay being primarily due to the posterior system. In this view, there is no reason to posit specialized neural systems whose functions are limited to those of short-term storage buffers. Prefrontal cortex provides the attentional pointer system for maintaining activation in the appropriate posterior processing systems. Short-term memory capacity and phenomena such as displacement of information in short-term memory are determined by limitations on the number of pointers that can be sustained by the prefrontal control systems. Key Words: coherence; event-related potentials; imaging; long-term memory; memory; short-term memory; working memory.
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Ruchkin et al. ascribe a pivotal role to long-term memory representations and binding within working memory. Here we focus on the interaction of working memory and long-term memory in supporting on-line representations of experience available to guide on-going processing, and we distinguish the role of frontal-lobe systems from what the hippocampus contributes to relational long-term memory binding.
The identity of working-memory and long-term memory representations follows from many lines of evidence. However, the data provided by Ruchkin et al. are hardly compelling, as they make unproved assumptions about hypothetical generators. We cite studies from our lab in which congruent slow-wave topographies were found for short-term and long-term memory tasks, strongly suggesting that both activate identical cell assemblies.
We focus on the functional specificity of theta and alpha oscillations and show that theta is related to working memory, whereas alpha is related to semantic long-term memory. Recent studies, however, indicate that alpha oscillations also play an important role during short-term memory retention and retrieval. This latter finding provides support for the basic hypothesis suggested by Ruchkin et al.
The goal of our target article is to establish that electrophysiological data constrain models of short-term memory retention operations to schemes in which activated long-term memory is its representational basis. The temporary stores correspond to neural circuits involved in the perception and subsequent processing of the relevant information, and do not involve specialized neural circuits dedicated to the temporary holding of information outside of those embedded in long-term memory. The commentaries ranged from general agreement with the view that short-term memory stores correspond to activated long-term memory (e.g., Abry, Sato, Schwartz, Loevenbruck & Cathiard [Abry etal.], Cowan, Fuster, Grote, Hickok & Buchsbaum, Keenan, Hyönä & Kaakinen [Keenan et al.], Martin, Morra), to taking a definite exception to this view (e.g., Baddeley, Düzel, Logie & Della Sala, Kroger, Majerus, Van der Linden, Colette & Salmon [Majerus et al.], Vallar).
Neural models have proposed how short-term memory (STM) storage in working memory and long-term memory (LTM) storage and recall are linked and interact, but are realized by different mechanisms that obey different laws. The authors' data can be understood in the light of these models, which suggest that the authors may have gone too far in obscuring the differences between these processes.
We challenge Ruchkin et al.'s claim in reducing short-term memory (STM) to the active part of long-term memory (LTM), by showing that their data cannot rule out the possibility that activation of posterior brain regions could also reflect the contribution of a verbal STM buffer.
The current model, based on event-related potential (ERP) studies, posits that the working-memory system is a state of activated long-term memory; this appears comprehensive, but it needs further detailed analysis of functional neural connectivity analysis within the prefrontal cortex (PFC) and between the posterior and prefrontal cortex. Specifically, the role of dorsolateral PFC and anterior cingulate cortex (ACC) is probably critical for PFC's attentional controller. Neural implementation of the executive function in working memory appears critical to build a firm model.
Single-unit data from the cortex of monkeys performing working-memory tasks support the main point of the target article. Those data, however, also indicate that the activation of long-term memory is essential to the processing of all cognitive functions. The activation of cortical long-term memory networks is a key neural mechanism in attention (working memory is a form thereof), perception, memory acquisition and retrieval, intelligence, and language.
Working-memory retention as activated long-term memory fails to capture orchestrated processing and storage, the hallmark of the concept of working memory. The event-related potential (ERP) data are compatible with working memory as a mental workspace that holds and manipulates information on line, which is distinct from long-term memory, and deals with the products of activated traces from stored knowledge.
Discussion of Daniel S. Ruchkin , Jordan Grafman , Katherine Cameron & Rita S. Berndt, Working memory retention systems: A state of activated long-term memory
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