The functions of sleep: A cognitive neuroscience perspective.

This Special Feature explores the various purposes served by sleep, describing current attempts to understand how the many functions of sleep are instantiated in neural circuits and cognitive structures. Our feature reflects current experts' opinions about, and insights into, the dynamic processes of sleep. In the last few decades, technological advances have supported the updated view that sleep plays an active role in both cognition and health. However, these roles are far from understood. This collection of articles evaluates the dynamic nature of sleep, how it evolves across the lifespan, becomes a competitive arena for memory systems through the influence of the autonomic system, supports the consolidation and integration of new memories, and how lucid dreams might originate. This set of papers highlights new approaches and insights that will lay the groundwork to eventually understand the full range of functions supported by sleep.

Sleep preferentially consolidates negative aspects of human memory: Well-powered evidence from two large online experiments.

Research suggests that sleep benefits memory. Moreover, it is often claimed that sleep selectively benefits memory for emotionally salient information over neutral information. However, not all scientists are convinced by this relationship [e.g., J. M. Siegel. Curr. Sleep Med. Rep., 7, 15-18 (2021)]. One criticism of the overall sleep and memory literature-like other literature-is that many studies are underpowered and lacking in generalizability [M. J. Cordi, B. Rasch. Curr. Opin. Neurobiol., 67, 1-7 (2021)], thus leaving the evidence mixed and confusing to interpret. Because large replication studies are sorely needed, we recruited over 250 participants spanning various age ranges and backgrounds in an effort to confirm sleep's preferential emotional memory consolidation benefit using a well-established task. We found that sleep selectively benefits memory for negative emotional objects at the expense of their paired neutral backgrounds, confirming our prior work and clearly demonstrating a role for sleep in emotional memory formation. In a second experiment also using a large sample, we examined whether this effect generalized to positive emotional memory. We found that while participants demonstrated better memory for positive objects compared to their neutral backgrounds, sleep did not modulate this effect. This research provides strong support for a sleep-specific benefit on memory consolidation for specifically negative information and more broadly affirms the benefit of sleep for cognition.

The effects of shared, depression-specific, and anxiety-specific internalizing symptoms on negative and neutral episodic memories following post-learning sleep.

Emotional memory bias is a common characteristic of internalizing symptomatology and is enhanced during sleep. The current study employs bifactor S-1 modeling to disentangle depression-specific anhedonia, anxiety-specific anxious arousal, and the common internalizing factor, general distress, and test whether these internalizing symptoms interact with sleep to influence memory for emotional and neutral information. Healthy adults (N = 281) encoded scenes featuring either negative objects (e.g., a vicious looking snake) or neutral objects (e.g., a chipmunk) placed on neutral backgrounds (e.g., an outdoor scene). After a 12-hour period of daytime wakefulness (n = 140) or nocturnal sleep (n = 141), participants judged whether objects and backgrounds were the same, similar, or new compared with what they viewed during encoding. Participants also completed the mini version of the Mood and Anxiety Symptom Questionnaire. Higher anxious arousal predicted worse memory across all stimuli features, but only after a day spent being awake-not following a night of sleep. No significant effects were found for general distress and anhedonia in either the sleep or wake condition. In this study, internalizing symptoms were not associated with enhanced emotional memory. Instead, memory performance specifically in individuals with higher anxious arousal was impaired overall, regardless of emotional valence, but this was only the case when the retention interval spanned wakefulness (i.e., not when it spanned sleep). This suggests that sleep may confer a protective effect on general memory impairments associated with anxiety.

The influence of encoding strategy on associative memory consolidation across wake and sleep.

Sleep benefits memory consolidation. However, factors present at initial encoding may moderate this effect. Here, we examined the role that encoding strategy plays in subsequent memory consolidation during sleep. Eighty-nine participants encoded pairs of words using two different strategies. Each participant encoded half of the word pairs using an integrative visualization technique, where the two items were imagined in an integrated scene. The other half were encoded nonintegratively, with each word pair item visualized separately. Memory was tested before and after a period of nocturnal sleep (N = 47) or daytime wake (N = 42) via cued recall tests. Immediate memory performance was significantly better for word pairs encoded using the integrative strategy compared with the nonintegrative strategy (P < 0.001). When looking at the change in recall across the delay, there was significantly less forgetting of integrated word pairs across a night of sleep compared with a day spent awake (P < 0.001), with no significant difference in the nonintegrated pairs (P = 0.19). This finding was driven by more forgetting of integrated compared with not-integrated pairs across the wake delay (P < 0.001), whereas forgetting was equivalent across the sleep delay (P = 0.26). Together, these results show that the strategy engaged in during encoding impacts both the immediate retention of memories and their subsequent consolidation across sleep and wake intervals.

The effects of cognitive reappraisal and sleep on emotional memory formation.

Emotion regulation (i.e. either up- or down-regulating affective responses to emotional stimuli) has been shown to modulate long-term emotional memory formation. Further, research has demonstrated that the emotional aspects of scenes are preferentially remembered relative to neutral aspects (known as the emotional memory trade-off effect). This trade-off is often enhanced when sleep follows learning, compared to an equivalent period of time spent awake. However, the interactive effects of sleep and emotion regulation on emotional memory are poorly understood. We presented 87 participants with pictures of neutral or negative objects on neutral backgrounds paired with instructions to either increase or decrease their emotional response by altering personal relevance, or to passively view the stimuli. Following a 12 h period of sleep or wakefulness, participants were tested for their memory of objects and backgrounds separately. Although we replicated the emotional memory trade-off effect, no differences in the magnitude of the trade-off effect were observed between regulation conditions. Sleep improved all aspects of memory, but it did not preferentially benefit memory for emotional components of scenes. Irrespective of a period of sleep or wake following encoding, findings suggest emotion regulation during encoding did not influence memory for emotional items at a 12-hour delay.

Sleep Spindles Preferentially Consolidate Weakly Encoded Memories.

Sleep has been shown to be critical for memory consolidation, with some research suggesting that certain memories are prioritized for consolidation. Initial strength of a memory appears to be an important boundary condition in determining which memories are consolidated during sleep. However, the role of consolidation-mediating oscillations, such as sleep spindles and slow oscillations, in this preferential consolidation has not been explored. Here, 54 human participants (76% female) studied pairs of words to three distinct encoding strengths, with recall being tested immediately following learning and again 6 h later. Thirty-six had a 2 h nap opportunity following learning, while the remaining 18 remained awake throughout. Results showed that, across 6 h awake, weakly encoded memories deteriorated the fastest. In the nap group, however, this effect was attenuated, with forgetting rates equivalent across encoding strengths. Within the nap group, consolidation of weakly encoded items was associated with fast sleep spindle density during non-rapid eye movement sleep. Moreover, sleep spindles that were coupled to slow oscillations predicted the consolidation of weak memories independently of uncoupled sleep spindles. These relationships were unique to weakly encoded items, with spindles not correlating with memory for intermediate or strong items. This suggests that sleep spindles facilitate memory consolidation, guided in part by memory strength.SIGNIFICANCE STATEMENT Given the countless pieces of information we encode each day, how does the brain select which memories to commit to long-term storage? Sleep is known to aid in memory consolidation, and it appears that certain memories are prioritized to receive this benefit. Here, we found that, compared with staying awake, sleep was associated with better memory for weakly encoded information. This suggests that sleep helps attenuate the forgetting of weak memory traces. Fast sleep spindles, a hallmark oscillation of non-rapid eye movement sleep, mediate consolidation processes. We extend this to show that fast spindles were uniquely associated with the consolidation of weakly encoded memories. This provides new evidence for preferential sleep-based consolidation and elucidates a physiological correlate of this benefit.

Medial prefrontal cortex has a causal role in selectively enhanced consolidation of emotional memories after a 24-hour delay: A TBS study.

Previous research points to an association between retrieval-related activity in the medial prefrontal cortex (mPFC) and preservation of emotional information compared to co-occurring neutral information following sleep. Although the role of the mPFC in emotional memory likely begins at encoding, little research has examined how mPFC activity during encoding interacts with consolidation processes to enhance emotional memory. This issue was addressed in the present study using transcranial magnetic stimulation in conjunction with an emotional memory paradigm. Healthy young adults encoded negative and neutral scenes while undergoing concurrent TMS with a modified short intermittent theta burst stimulation (sTBS) protocol. Participants received stimulation to either the mPFC or an active control site (motor cortex) during the encoding phase. Recognition memory for scene components (objects and backgrounds) was assessed after a short (30-minute) and a long delay (24-hour, including a night of sleep) to obtain measures of specific and gist-based memory processes. The results demonstrated that, relative to control stimulation, sTBS to the mPFC enhanced memory for negative objects on the long delay test (collapsed across specific and gist-based memory measures). mPFC stimulation had no discernable effect on memory for objects on the short delay test nor on the background images at either test. These results suggest that mPFC activity occurring during encoding interacts with consolidation processes to preferentially preserve negatively salient information.SIGNIFICANCE STATEMENT:Understanding how emotional information is remembered over time is critical to understanding memory in the real world. The present study used noninvasive brain stimulation (repetitive transcranial magnetic stimulation, rTMS) to investigate the interplay between mPFC activity that occurs during memory encoding and its subsequent interactions with consolidation processes. rTMS delivered to the mPFC during encoding enhanced memory for negatively valenced pictures on a test following a 24-hr delay, with no such effect on a test occurring shortly after the encoding phase. These results are consistent with the hypothesis that emotional aspects of memories are differentially subjected to consolidation processes, and that the mPFC might contribute to this "tag-and-capture" mechanism during the initial formation of such memories.

Slow oscillation-spindle coupling is negatively associated with emotional memory formation following stress.

Both stress and sleep enhance emotional memory. They also interact, with the largest effect of sleep on emotional memory being seen when stress occurs shortly before or after encoding. Slow wave sleep (SWS) is critical for long-term episodic memory, facilitated by the temporal coupling of slow oscillations and sleep spindles. Prior work in humans has shown these associations for neutral information in non-stressed participants. Whether coupling interacts with stress to facilitate emotional memory formation is unknown. Here, we addressed this question by reanalyzing an existing dataset of 64 individuals. Participants underwent a psychosocial stressor (32) or comparable control (32) prior to the encoding of 150-line drawings of neutral, positive, and negative images. All participants slept overnight with polysomnography, before being given a surprise memory test the following day. In the stress group, time spent in SWS was positively correlated with memory for images of all valences. Results were driven by those who showed a high cortisol response to the stressor, compared to low responders. The amount of slow oscillation-spindle coupling during SWS was negatively associated with neutral and emotional memory in the stress group only. The association with emotional memory was significantly stronger than for neutral memory within the stress group. These results suggest that stress around the time of initial memory formation impacts the relationship between slow wave sleep and memory.

Distinct stress-related changes in intrinsic amygdala connectivity predict subsequent positive and negative memory performance.

Experiencing stress before an event can influence how that event is later remembered. In the current study, we examine how individual differences in one's physiological response to a stressor are related to changes to underlying brain states and memory performance. Specifically, we examined how changes in intrinsic amygdala connectivity relate to positive and negative memory performance as a function of stress response, defined as a change in cortisol. Twenty-five participants underwent a social stressor before an incidental emotional memory encoding task. Cortisol samples were obtained before and after the stressor to measure individual differences in stress response. Three resting state scans (pre-stressor, post-stressor/pre-encoding and post-encoding) were conducted to evaluate pre- to post-stressor and pre- to post-encoding changes to intrinsic amygdala connectivity. Analyses examined relations between greater cortisol changes and connectivity changes. Greater cortisol increases were associated with a greater decrease in prefrontal-amygdala connectivity following the stressor and a reversal in the relation between prefrontal-amygdala connectivity and negative vs. positive memory performance. Greater cortisol increases were also associated with a greater increase in amygdala connectivity with a number of posterior sensory regions following encoding. Consistent with prior findings in non-stressed individuals, pre- to post-encoding increases in amygdala-posterior connectivity were associated with greater negative relative to positive memory performance, although this was specific to lateral rather than medial posterior regions and to participants with the greatest cortisol changes. These findings suggest that stress response is associated with changes in intrinsic connectivity that have downstream effects on the valence of remembered emotional content.

Memory for emotional images across sleep versus wake in school-aged children.

Sleep is important for emotional well-being, memory, and development in children. Regarding memory, sleep has been shown to advantage accuracy for declarative tasks but not procedural tasks. There is some evidence to suggest that sleep provides a relatively greater benefit for memory of negative emotional versus neutral images. However, the extent to which sleep benefits emotionally positive memories in children is not clear. This study assessed memory after nocturnal sleep versus daytime wake in a within-person design involving a sample of 40 children aged 7 to 14 years (M = 10.6 ± 1.9 years; 18 boys and 22 girls) for images of negative, neutral, and positive valences. Results show that after accounting for response time, memory accuracy overall was greater after sleep compared with equivalent time of wake and memory accuracy was greatest for positive images compared with both negative and neutral images. However, there was no difference between memory for negative images and that for neutral images in children, and there was no condition by valence interaction. Sleep characteristics as recorded using actigraphy, diary, and parent report were not predictive of memory performance after sleep when correcting for multiple comparisons. Overall, the results suggest that sleep may benefit memory in otherwise healthy children but that despite a bias toward memory for positive items over both negative and neutral items, there is not a relatively greater benefit for emotional versus neutral memory consolidation across sleep periods compared with wake periods.

Sleep spectral power correlates of prospective memory maintenance.

Prospective memory involves setting an intention to act that is maintained over time and executed when appropriate. Slow wave sleep (SWS) has been implicated in maintaining prospective memories, although which SWS oscillations most benefit this memory type remains unclear. Here, we investigated SWS spectral power correlates of prospective memory. Healthy young adult participants completed three ongoing tasks in the morning or evening. They were then given the prospective memory instruction to remember to press "Q" when viewing the words "horse" or "table" when repeating the ongoing task after a 12-h delay including overnight, polysomnographically recorded sleep or continued daytime wakefulness. Spectral power analysis was performed on recorded sleep EEG. Two additional groups were tested in the morning or evening only, serving as time-of-day controls. Participants who slept demonstrated superior prospective memory compared with those who remained awake, an effect not attributable to time-of-day of testing. Contrary to prior work, prospective memory was negatively associated with SWS. Furthermore, significant increases in spectral power in the delta-theta frequency range (1.56 Hz-6.84 Hz) during SWS was observed in participants who failed to execute the prospective memory instructions. Although sleep benefits prospective memory maintenance, this benefit may be compromised if SWS is enriched with delta-theta activity.

Higher post-encoding cortisol benefits the selective consolidation of emotional aspects of memory.

Emotional experiences create durable memory traces in the brain, especially when these memories are consolidated in the presence of stress hormones such as cortisol. Although some research suggests cortisol elevation can increase long-term memory for emotional relative to neutral content, the impact of stress and cortisol on the consolidation of emotional and neutral aspects of memories when they are part of the same experience remains unknown. Here, after encoding complex scenes consisting of negative or neutral objects placed on neutral backgrounds, participants were exposed to a psychosocial stressor (or matched control condition) in order to examine the impact of stress and cortisol on early consolidation processes. The next day, once cortisol levels had returned to baseline, specific and gist recognition memory were tested separately for objects and backgrounds. Results indicate that while there was a numerical increase in memory for negative objects in the stress group, higher endogenous cortisol concentrations were specifically associated with decreased memory for the neutral backgrounds originally paired with negative objects. Moreover, across all participants, cortisol levels were positively correlated with the magnitude of the emotional memory trade-off effect. Specifically, while memory for negative objects was preserved, elevated cortisol during early consolidation was associated with decreased memory for neutral backgrounds that were initially paired with negative objects. These memory effects were observed in both the stricter specific measure of memory and the less conservative measure of gist memory. Together, these findings suggest that rather than influencing all aspects of an experience similarly, elevated cortisol during early consolidation selectively preserves what is most emotionally salient and adaptive to remember while allowing the loss of memory for less important neutral information over time.

Interactive effects of stress reactivity and rapid eye movement sleep theta activity on emotional memory formation.

Sleep and stress independently enhance emotional memory consolidation. In particular, theta oscillations (4-7 Hz) during rapid eye movement (REM) sleep increase coherence in an emotional memory network (i.e., hippocampus, amygdala, and prefrontal cortex) and enhance emotional memory. However, little is known about how stress during learning might interact with subsequent REM theta activity to affect emotional memory. In the current study, we examined whether the relationship between REM theta activity and emotional memory differs as a function of pre-encoding stress exposure and reactivity. Participants underwent a psychosocial stressor (the Trier Social Stress Task; n = 32) or a comparable control task (n = 32) prior to encoding. Task-evoked cortisol reactivity was assessed by salivary cortisol rise from pre- to post-stressor, and participants in the stress condition were additionally categorized as high or low cortisol responders via a median split. During incidental encoding, participants studied 150 line drawings of negative, neutral, and positive images, followed by the complete color photo. All participants then slept overnight in the lab with polysomnographic recording. The next day, they were given a surprise recognition memory task. Results showed that memory was better for emotional relative to neutral information. Critically, these findings were observed only in the stress condition. No emotional memory benefit was observed in the control condition. In stressed participants, REM theta power significantly predicted memory for emotional information, specifically for positive items. This relationship was observed only in high cortisol responders. For low responders and controls, there was no relationship between REM theta and memory of any valence. These findings provide evidence that elevated stress at encoding, and accompanying changes in neuromodulators such as cortisol, may interact with theta activity during REM sleep to promote selective consolidation of emotional information.

Acute sleep deprivation and the selective consolidation of emotional memories.

Research suggests that sleep preferentially consolidates the negative aspects of memories at the expense of the neutral aspects. However, the mechanisms by which sleep facilitates this emotional memory trade-off remain unknown. Although active processes associated with sleep-dependent memory consolidation have been proposed to underlie this effect, this trade-off may also be modulated by non-sleep-related processes, such as the circadian factors, stress-related factors, and/or mood congruent context effects involved in sleep deprivation. We sought to examine the potential role of these factors by randomizing 39 young adults into either a total sleep deprivation condition (26 consecutive hours awake) or a sleep condition (8 h sleep opportunity). Replicating the emotional memory trade-off effect, negative objects were better remembered than neutral objects or background images. However, in spite of generally worse memory performance (for neutral and background information), sleep-deprived participants showed similar recognition rates for negative emotional memories relative to participants who were given a full night of sleep.

Overnight sleep benefits both neutral and negative direct associative and relational memory.

Strong evidence suggests that sleep plays a role in memory consolidation, which involves both stabilizing memory into long-term storage as well as integrating new information into existing stores. The current study investigated consolidation, across a day of wakefulness or night of sleep, of emotional and neutral directly learned visual paired associates (A-B/B-C pairs) as well as formation of memory for relational pairs formed via overlapping learned components (A-C pairs). Participants learned 40 negative and 40 neutral face-object pairs followed by a baseline test in session 1 either in the morning or evening. They then spent a 12-hour retention period during which participants either went about their normal day or spent the night in the sleep lab. During session 2, participants completed a surprise test to assess their memory for relational pairs (A-C) as well as memory for direct associates (A-B/B-C). As hypothesized, the results demonstrated that a 12-hour retention period predominantly spent asleep, compared to awake, benefited memory for both relational and direct associative memory. However, contrary to the hypothesis that emotional salience would promote preferential consolidation, sleep appeared to benefit both negative and neutral information similarly for direct associative and relational memories, suggesting that sleep may interact with other factors affecting encoding (e.g., depth of encoding) to benefit direct and relational associative memory. As one of the few studies examining the role of nocturnal sleep and emotion on both direct and relational associative memory, our findings suggest key insights into how overnight sleep consolidates these different forms of memory.

Do different salience cues compete for dominance in memory over a daytime nap?

Information that is the most salient and important for future use is preferentially preserved through active processing during sleep. Emotional salience is a biologically adaptive cue that influences episodic memory processing through interactions between amygdalar and hippocampal activity. However, other cues that influence the importance of information, such as the explicit direction to remember or forget, interact with the inherent salience of information to determine its fate in memory. It is unknown how sleep-based processes selectively consolidate this complex information. The current study examined the development of memory for emotional and neutral information that was either cued to-be-remembered (TBR) or to-be-forgotten (TBF) across a daytime period including either napping or wakefulness. Baseline memory revealed dominance of the TBR cue, regardless of emotional salience. As anticipated, napping was found to preserve memory overall significantly better than remaining awake. Furthermore, we observed a trending interaction indicating that napping specifically enhanced the discrimination between the most salient information (negative TBR items) over other information. We found that memory for negative items was positively associated with the percentage of SWS obtained during a nap. Furthermore, the magnitude of the difference in memory between negative TBR items and negative TBF items increased with greater sleep spindle activity. Taken together, our results suggest that although the cue to actively remember or intentionally forget initially wins out, active processes during sleep facilitate the competition between salience cues to promote the most salient information in memory.

Post-encoding stress enhances mnemonic discrimination of negative stimuli.

Stress influences how we remember emotional events and how these events shape future behaviors. However, the impact of stress on memory specificity for emotional events has yet to be examined. To this end, the present study utilized a mnemonic discrimination task that taxes hippocampal pattern separation, the process of distinguishing between overlapping experiences, thereby allowing us to better understand the mechanisms by which stress affects gist versus detail memory of emotional events. Participants encoded scenes composed of negative or neutral objects placed on neutral backgrounds and then underwent a psychosocial stressor or matched control task. Twenty-four hours later during testing, objects were presented separately, with some identical old objects (targets), some new objects (foils), and some similar but not identical objects (lures). Target recognition was enhanced for negative compared to neutral objects in both the stress and control groups. Interestingly, post-encoding stress selectively enhanced mnemonic discrimination of negative versus neutral objects, which was not the case in the control group. Measures of salivary cortisol revealed a quadratic inverted U relationship between negative mnemonic discrimination and cortisol increase. These findings suggest that moderate cortisol release following stress is associated with enhanced memory precision for negative information.

Residual effects of emotion are reflected in enhanced visual activity after sleep.

Research has investigated how sleep affects emotional memory and how emotion enhances visual processing, but these questions are typically asked by re-presenting an emotional stimulus at retrieval. For the first time, we investigate whether sleep affects neural activity during retrieval when the memory cue is a neutral context that was previously presented with either emotional or nonemotional content during encoding. Participants encoded scenes composed of a negative or neutral object on a neutral background either in the morning (preceding 12 hours awake; wake group) or evening (preceding 12 hours including a night of sleep; sleep group). At retrieval, participants viewed the backgrounds without their objects, distinguishing new backgrounds from those previously studied. Occipital activity was greater within the sleep group than the wake group specifically during the successful retrieval of neutral backgrounds that had been studied with negative (but not neutral) objects. Moreover, there was enhanced connectivity between the middle occipital gyrus and hippocampus following sleep. Within the sleep group, the percentage of REM sleep obtained correlated with activity in the middle occipital gyrus, lingual gyrus, and cuneus during the successful retrieval of neutral backgrounds previously paired with negative objects. These results confirm that emotion affects neural activity during retrieval even when the cues themselves are neutral, and demonstrate, for the first time, that this residual effect of emotion on visual activity is greater after sleep and may be maximized by REM sleep.

The impact of sleep on true and false memory across long delays.

While the influence of sleep on memory has a long history, sleep's role in the formation of false memories is less clear. Moreover, virtually nothing is known about the development of false memories beyond delays of about 12h. Here, for the first time, we assess post-sleep development of true and false memories across longer delay intervals of 24 and 48h. Although technically a false memory, remembering information that is related to the theme, or gist, of an experience can be considered an adaptive process. Some evidence suggests that sleep, compared to a wake period, increases both true and gist-based false memories in the Deese-Roediger-McDermott (DRM) task, but not all studies have returned this result, and most studies cannot rule out the possibility that sleep is merely protecting the information from interference, as opposed to actively aiding its consolidation. Here, to equate amount of time spent awake and asleep across groups, we assess how the positioning of sleep relative to memory encoding impacts retention across longer delays of 24 and 48h. Participants encoded 16 DRM lists in the morning (WAKE 1st Groups) or evening (SLEEP 1st Groups), and were tested either 24 or 48h later at the same time of day. Results demonstrate that true memory is better when participants sleep soon after learning. Sleeping first also increased false memory, but only in low performers. Importantly, and similar to previous studies, we found a negative correlation between slow-wave sleep (SWS) and false memory, suggesting that SWS may be detrimental for semantic/gist processing.

The effects of sleep restriction and sleep deprivation in producing false memories.

False memory has been claimed to be the result of an associative process of generalisation, as well as to be representative of memory errors. These can occur at any stage of memory encoding, consolidation, or retrieval, albeit through varied mechanisms. The aim of this paper is to experimentally determine: (i) if cognitive dysfunction brought about by sleep loss at the time of stimulus encoding can influence false memory production; and (ii) whether this relationship holds across sensory modalities. Subjects undertook both the Deese-Roedigger-McDermott (DRM) false memory task and a visual task designed to produce false memories. Performance was measured while subjects were well-rested (9h Time in Bed or TIB), and then again when subjects were either sleep restricted (4h TIB for 4 nights) or sleep deprived (30h total SD). Results indicate (1) that partial and total sleep loss produced equivalent effects in terms of false and veridical verbal memory, (2) that subjects performed worse after sleep loss (regardless of whether this was partial or total sleep loss) on cued recognition-based false and veridical verbal memory tasks, and that sleep loss interfered with subjects' ability to recall veridical, but not false memories under free recall conditions, and (3) that there were no effects of sleep loss on a visual false memory task. This is argued to represent the dysfunction and slow repair of an online verbal associative process in the brain following inadequate sleep.