This article investigates several consequences of a recent trend in philosophy of mind to shift the relata of realization from mental state–physical state to function‐mechanism. It is shown, by applying both frameworks to the neuroscientific case study of memory consolidation, that, although this shift can be used to avoid the immediate antireductionist consequences of the traditional argument from multiple realizability, what is gained is a far more modest form of reductionism than recent philosophical accounts have intimated and neuroscientists (...) themselves have claimed. (shrink)

This paper argues that the biochemistry of memory consolidation provides valuable model systems for exploring the multiple realization of psychological states.

Based on brain state-dependent behavioral changes, consolidation of sensorimotor memories has been posited to evolve in two different functional stages. Only the second of these stages requires sleep and leads to performance benefits. Recent results, however, suggest that sleep is not always crucial for the expression of delayed behavioral gains but might be critical for enhancing automaticity in the absence of attention, another expression of memory consolidation.

Following Karni's seminal work, Walker and other researchers have recently provided gradually convincing evidence that sleep is critical for the consolidation-based enhancement (CBE) of motor sequence learning. Studies in our laboratory using a motor adaptation paradigm, however, show that CBE can also occur after the simple passage of time, suggesting that sleep effects on memory consolidation are task-related, and possibly dependent on anatomically dissociable circuits.

Does sleep restore brain function or does it consolidate memory? I argue that memory consolidation during sleep is an offshoot of restitution. Continual learning causes local synapse-specific neural fatigue, which then masks expression of that learning, especially on time-limited tests of procedural skills. Sleep serves to restore the fatigued synapses, revealing the consolidation-based enhancement observed as a “latent” overnight improvement in learning.

We present evidence disputing the hypothesis that memories are processed or consolidated in REM sleep. A review of REM deprivation (REMD) studies in animals shows these reports to be about equally divided in showing that REMD does, or does not, disrupt learning/memory. The studies supporting a relationship between REM sleep and memory have been strongly criticized for the confounding effects of very stressful REM deprivation techniques. The three major classes of antidepressant drugs, monoamine oxidase inhibitors (MAOIs), tricyclic antidepressants (...) (TCAs), and selective serotonin reuptake inhibitors (SSRIs), profoundly suppress REM sleep. The MAOIs virtually abolish REM sleep, and the TCAs and SSRIs have been shown to produce immediate (40–85%) and sustained (30–50%) reductions in REM sleep. Despite marked suppression of REM sleep, these classes of antidepressants on the whole do not disrupt learning/memory. There have been a few reports of patients who have survived bilateral lesions of the pons with few lingering complications. Although these lesions essentially abolished REM sleep, the patients reportedly led normal lives. Recent functional imaging studies in humans have revealed patterns of brain activity in REM sleep that are consistent with dream processes but not with memory consolidation. We propose that the primary function of REM sleep is to provide periodic endogenous stimulation to the brain which serves to maintain requisite levels of central nervous system (CNS) activity throughout sleep. REM is the mechanism used by the brain to promote recovery from sleep. We believe that the cumulative evidence indicates that REM sleep serves no role in the processing or consolidation of memory. Key Words: antidepressant drugs; brain stem lesions; dreams; functional imaging; memory consolidation; REM deprivation; REM sleep; theta rhythm. (shrink)

This study examined the effect of an interference task on the consolidation of duration in long-term memory. In a temporal generalization task, the participants performed a learning phase with a reference duration that either was, or was not, followed 30 minutes later by a 15-min interference task. They were then given a memory test, 24h later. Using different participant groups, several reference durations were examined, from several hundred milliseconds (600ms) to several seconds (2.5, 4 and 8s). The (...) results showed that the scalar timing property (i.e. precision proportional to judged duration) was preserved despite the interference task given during the memory consolidation process. However, the interference task increased the variability of time judgment and tended to produce a lengthening effect in all reference duration conditions. The modeling of individual data with parameters derived from scalar expectancy theory suggests that disrupting the memory consolidation of learned reference durations introduces noise in their representation in memory, with time being specifically distorted towards a lengthened duration. (shrink)

Clinical and experimental evidence suggests that hippocampal damage causes more severe disruption of episodic memories if those memories were encoded in the recent rather than the more distant past. This decrease in sensitivity to damage over time might reflect the formation of multiple traces within the hippocampus itself, or the formation of additional associative links in entorhinal and association cortices. Physiological evidence also supports a two-stage model of the encoding process in which the initial encoding occurs during active waking and (...) deeper consolidation occurs via the formation of additional memory traces during quiet waking or slow-wave sleep. In this article I will describe the changes in cholinergic tone within the hippocampus in different stages of the sleep-wake cycle and will propose that these changes modulate different stages of memory formation. In particular, I will suggest that the high levels of acetylcholine that are present during active waking might set the appropriate dynamics for encoding new information in the hippocampus, by partially suppressing excitatory feedback connections and so facilitating encoding without interference from previously stored information. By contrast, the lower levels of acetylcholine that are present during quiet waking and slow-wave sleep might release this suppression and thereby allow a stronger spread of activity within the hippocampus itself and from the hippocampus to the entorhinal cortex, thus facilitating the process of consolidation of separate memory traces. (shrink)

Neural reuse posits development of functional overlap in brain system circuits to accommodate complex evolutionary functions. Evolutionary adaptation evolved neural circuits that have been exploited for many uses. One such use is engaging cognitive processes in memory consolidation during the neurobiological states of sleep. Neural reuse, therefore, should not be limited to neural circuitry, but be extended to include sleep-state associated memory processes.

Unfortunately, some researchers think a good scientific theory is one that has been repeatedly confirmed, and a bad theory is one that has not received consistent confirmation. However, confirmation of a theory depends on the extent to which a hypothesis exposes itself to disconfirmation. One confirmation of a highly specific, falsifiable experiment can have a far greater impact than the disconfirmation of twenty experiments that are virtually unfalsifiable. This commentary (1) counteracts misleading biases regarding the REM sleep/memory consolidation (...) theory, and (2) demonstrates how chaotic cerebral activation during sleep is an essential component of long-term memory storage processes. [Vertes & Eastman]. (shrink)

Visuomotor adaptation to novel environments can occur via non-physical means, such as observation. Observation does not appear to activate the same implicit learning processes as physical practice, rather it appears to be more strategic in nature. However, there is evidence that interspersing observational practice with physical practice can benefit performance and memory consolidation either through the combined benefits of separate processes or through a change in processes activated during observation trials. To test these ideas, we asked people to (...) practice aiming to targets with visually rotated cursor feedback or engage in a combined practice schedule comprising physical practice and observation of projected videos showing successful aiming. Ninety-three participants were randomly assigned to one of five groups: massed physical practice, distributed physical practice, or one of 3 types of combined practice: alternating blocks, or all observation before or after blocked physical practice. Participants received 100 practice trials. All groups improved in adaptation trials and showed savings across the 24-h retention interval relative to initial practice. There was some forgetting for all groups, but the magnitudes were larger for physical practice groups. The Act and Obs_During groups were most accurate in retention and did not differ, suggesting that observation can serve as a replacement for physical practice if supplied intermittently and offers advantages above just resting. However, after-effects associated with combined practice were smaller than those for physical practice control groups, suggesting that beneficial learning effects as a result of observation were not due to activation of implicit learning processes. Reaction time, variable error, and post-test rotation drawings supported this conclusion that adaptation for observation groups was promoted by explicit/strategic processes. (shrink)

Memory consolidation is the process in which memory traces are strengthened over time for later retrieval. Although some theories hold that consolidation can only occur during sleep, accumulating evidence suggests that brief periods of wakeful rest may also facilitate consolidation. Interestingly, however, Varma and colleagues reported that a demanding 2-back task following encoding produced a similar performance to a wakeful reset condition. We tested whether participants’ recall would be best following a wakeful rest condition as (...) compared to other distractor conditions, consistent with the extant wakeful rest literature, or whether we would replicate the finding by Varma and colleagues such that participants’ memory benefitted from both a rest and a 2-back task following encoding. Across two experiments, we used similar and the same encoding material as used the one by Varma and colleagues, employed a wakeful rest condition adapted for online testing, and compared participants’ recall across post-encoding conditions. In the first experiment, we used a between-subjects design and compared participants’ cued recall performance following a period of wakeful rest, a 2-back task, or a rest + sounds condition. The second experiment more closely replicated the experimental design used by Varma and colleagues using a within-subjects manipulation. Ultimately, our findings more consistently aligned with the canonical wakeful rest finding, such that recall was better following the rest condition than all other post-encoding conditions. These results support the notion that wakeful rest may allow for consolidation by protecting recently encoded information from interference, thereby improving memory performance. (shrink)

Matthew Walker's model overcomes the unrefined classical concept of consolidation as a unitary process. Presently still confined in its scope to selective data mainly referring to procedural motor learning, the model nonetheless provides a valuable starting point for further refinements, which would be required for a more comprehensive account of different types and aspects of human memory consolidation.

AimRapid eye movement (REM) sleep, non-rapid eye movement (NREM) sleep, and sleep spindles are all implicated in the consolidation of procedural memories. Relative contributions of sleep stages and sleep spindles were previously shown to depend on individual differences in task processing. However, no studies to our knowledge have focused on individual differences in experience with Vipassana meditation as related to sleep. Vipassana meditation is a form of mental training that enhances proprioceptive and somatic awareness and alters attentional style. The (...) goal of this study was to examine a potential role for Vipassana meditation experience in sleep-dependent procedural memory consolidation.MethodsGroups of Vipassana meditation practitioners (N = 22) and matched meditation-naïve controls (N = 20) slept for a daytime nap in the laboratory. Before and after the nap they completed a procedural task on the Wii Fit balance platform.ResultsMeditators performed slightly better on the task before the nap, but the two groups improved similarly after sleep. The groups showed different patterns of sleep-dependent procedural memory consolidation: in meditators, task learning was positively correlated with density of slow occipital spindles, while in controls task improvement was positively associated with time in REM sleep. Sleep efficiency and sleep architecture did not differ between groups. Meditation practitioners, however, had a lower density of occipital slow sleep spindles tha... (shrink)

Review of Individual and Colllective Consolidation. Analogous Processes on Different Levels.

From its genesis in the 1960s, the focus of inquiry in neuroscience has been on the cellular and molecular processes underlying neural activity. In this pursuit neuroscience has been enormously successful. Like any successful scientific inquiry, initial successes have raised new questions that inspire ongoing research. While there is still much that is not known about the molecular processes in brains, a great deal of very important knowledge has been secured, especially in the last 50 years. It has also attracted (...) the attention of a number of philosophers, some of whom have viewed it as evidence for a ruthlessly reductionistic program that will eventually explain how mental processes are performed in the brain in purely molecular terms. As neuroscience developed, however, there emerged a smaller group of researchers who focused on systems, behavioral, and cognitive neuroscience. These investigators have also made impressive advances in the last 50 years and they have been the focus of an even larger group of philosophers, who have appealed to systems level understanding of the brain as providing the appropriate point of connection to the information processing accounts advanced in psychology. (shrink)

The function of REM, or any other stage of sleep, can currently only be conjectured. A rational evaluation of the role of REM in memory processing requires systematic testing of hypotheses that are optimally derived from a complete synthesis of existing knowledge. Our view is that the large number of studies supporting a relationship between REM-related brain activity and memory is not easily explained away. [Vertes & Eastman].

Is consolidation needed to account for retroactive interference in free recall? Interfering mental activity during the retention interval of a memory task impairs performance, in particular if the interference occurs in temporal proximity to the encoding of the to-be-remembered information. There are at least two rival theoretical accounts of this temporal gradient of retroactive interference. The cognitive neuroscience literature has suggested neural consolidation is a pivotal factor determining item recall. According to this account, interfering activity interrupts (...) class='Hi'>consolidation processes that would otherwise stabilize the memory representations of TBR items post-encoding. Temporal distinctiveness theory, by contrast, proposes that the retrievability of items depends on their isolation in psychological time. According to this theory, information processed after the encoding of TBR material will reduce the temporal distinctiveness of the TBR information. To test between these accounts, implementations of consolidation were added to the SIMPLE model of memory and learning. We report data from two experiments utilizing a two-list free recall paradigm. Modeling results imply that SIMPLE was able to model the data and did not benefit from the addition of consolidation. It is concluded that the temporal gradient of retroactive interference cannot be taken as evidence for memory consolidation. (shrink)

In this Hypothesis and Theory paper, we consider the problem of learning deeply structured knowledge representations in the absence of predefined ontologies, and in the context of long-term learning. In particular, we consider this process as a sequence of re-representation steps, of various kinds. The Information Dynamics of Thinking theory (IDyOT) admits such learning, and provides a hypothetical mechanism for the human-like construction of hierarchical memory, with the provision of symbols constructed by the system that embodies the theory. The (...) combination of long-term learning and meaning construction in terms of symbols grounded in perceptual experience entails that the system, like a human, be capable of memory consolidation, to manage the complex and inconsistent structures that can result from learning of information that becomes more complete over time. Such consolidation changes memory structures, and thus changes their meaning. Therefore, memory consolidation entails re-representation, while re-representation entails changes of meaning. Ultimately, the theory proposes that the processes of learning and consolidation should be considered as repeated re-representation of what is learned. (shrink)

We argue that by neglecting the fact that procedural memory may also have episodic qualities, and by considering only a systems approach to memory, Walker's account of consolidation of learning during subsequent sleep ignores alternative accounts of how sleep stages may be interdependent. We also question the proposition that sleep-based consolidation largely bypasses hippocampal structures.

Recent evidence from our lab indicates that LTP shares an important property with memory consolidation: it is consolidated by natural reinforcement. Nevertheless, the hypothesis, that LTP-like mechanisms or other forms of enhanced synaptic efficacy are basic elements in learning is not unequivocally supported. Skepticism aside, LTP is an accessible experimental model that is optimally equipped for the investigation of the cellular and molecular machinery involved in synaptic weight changes.

The progressive multistage stabilization of memory (consolidation) relies on post-acquisition neural reorganization. We hypothesize that two processes subserve procedural memory consolidation and are reflected in delayed post-acquisition performance gains: (1) synaptic consolidation, which is classical Hebbian, and (2) in some tasks, concurrently or consequently, “system consolidation,” which might in some skills be sleep-dependent. Behavioral interference may affect either type of consolidation.

Research on evaluative conditioning (EC) shows that attitudes can emerge from co-occurrences of stimuli, and accumulating evidence suggests that EC usually depends on memory for these stimulus contingencies. Therefore, processes known to aid memory retention may be relevant for the development of stable attitudes. One such process may be memory consolidation, assumed to be promoted by waking rest and sleep. In two pre-registered experiments, we investigated whether waking rest (vs. cognitive activity, Experiment 1) and sleep (vs. (...) wakefulness, Experiment 2) in between conditioning and measurement of EC, consolidate contingency memory and EC. Contrary to our predictions, waking rest (vs. cognitive activity) promoted neither contingency memory nor EC effects. Sleep (vs. wakefulness) decreased forgetting of contingency memory but crucially, it did not attenuate the impact of counterconditioning on contingency memory. Sleep also did not influence EC effects, nor the reduction of EC by counterconditioning. EC effects in both experiments were predicted by contingency memory. Yet, unexpectedly, EC effects occurred in the absence of contingency memory after waking rest, but neither after sleep nor in the active control conditions. Our findings emphasise a role of contingency memory in EC, but it remains unclear whether this role changes during waking rest. (shrink)

This paper aims to reconfigure the place of memory in epistemology. I start by rethinking the problem that memory systems solve; rather than merely functioning to store information, I argue that the core function of any memory system is to support accurate and relevant retrieval. This way of specifying the function of memory has consequences for which structures and mechanisms make up a memory system. In brief, memory systems are modeling systems. This means that (...) they generate, update and manage a series of overlapping, simplified, relational representations that map out features of the world. Succeeding at building and maintaining models requires the kind of active knowledge generation traditionally associated only with deliberative reasoning. (shrink)