Sleep and Memory: Retention 8 and 24 Hours After Initial Learning

Retention of nonsense syllables was tested in two groups of Ss: those having initial learning in the morning and those having initial learning at night immediately before sleep. These groups were subdivided so that recall was tested either 8 or 24 hrs after initial learning. As Jenkins and Dallenbach first demonstrated, retention was superior after 8 hrs for Ss when learned at night compared to those who learned in the morning. Retention with night learning was equal after 24 hrs to that observed after 8 hrs. Surprisingly, retention scores after morning learning were superior after 24 hrs to those observed after 8 hrs. Some possible interpretations of this result are advanced.     

Effects of sleep loss on waking actigraphy

STUDY OBJECTIVES: To assess the effect of sleep loss and the effect of a sedating drug on waking actigraphy DESIGN: N/A SETTING: N/A PARTICIPANTS: Seventeen healthy volunteers, aged 19-35 yrs Interventions: Four night-day treatments presented in a Latin Square Design: placebo-8 hr time-in-bed (TIB), placebo-4 hr TIB, placebo-0 hr TIB, and diphenhydramine 50 mg-8 hr TIB. MEASUREMENTS AND RESULTS: After the appropriate TIB, medication was administered at 09:00 hr, the Multiple Sleep Latency Test at 09:30, 11:30, 13:30, 15:30, and 17:30 hr, and a 45 min performance battery at 10:30, 14:30, and 16:30 hr. Each day the volunteers wore actigraphs from 0700-1800 hrs. Decreasing TIB was associated with decreased daily mean sleep latency on the MSLT with 4 and 0 hrs differing from 8 hrs and each other. Daytime activity also was reduced by the reduced prior TIB. Increased inactivity relative to the 8 hr TIB developed between the 4 hr and 0 hr TIBs, with 4 hrs differing from 0 hrs, but not 8 hrs. Diphenhydramine 50 mg reduced mean daily sleep latency and increased percent inactive time relative to placebo. On the MSLT diphenhydramine was intermediate to 4 hr and 0 hr TIB and on actigraphy it was similar to 0 hr TIB. CONCLUSIONS: The difference in the effect of diphenhydramine on these actigraphy and MSLT may reflect the different sensitivities of the measures.     

The sleep of conjoined twins.

Conjoined twins with a common heart system and circulatory system were observed during the 14th and 15th days for 11 hr. Sleep, Waking, and Quiet and Active Sleep were recorded in minute intervals. Clear independence of sleep and waking was manifest, and total independence of Quiet and Active sleep was noted.     

Duration and Placement of Sleep in a "Disentrained" Environment

Sleep/wake cycles of 9 young adults were electrographically recorded during 60 hrs of enforced bedrest. During this period subjects were required to lie quietly, with no time cues and minimal exogenous stimulation. Sleep and wakefulness patterns were clearly modified under these conditions. There was an alternation of waking periods with an average length of 2.7 hrs and sleep episodes with a mean duration of 2.99 hrs. Eighty percent of both sleep and waking periods were less than 4 hrs duration. The circadian pattern of sleep period duration persisted in disentrainment, but temporal organization of sleep episodes was substantially disrupted; sleep episodes occurred throughout the 24-hr day. The results suggest the presence of two distinct components of the human sleep system–one, sleep duration, is controlled by an endogenous circadian oscillatory system; another, sleep placement, is controlled primarily by behavioral controls, in the form of social and occupational pressures, and self-imposed behavioral alternatives to sleep.     

The characteristics of recovery sleep when recovery opportunity is restricted

STUDY OBJECTIVES: The aim of this study was to investigate the recovery of sleep and waking functions following one night of total sleep deprivation, when recovery opportunity was either augmented or restricted. DESIGN: The 9-day laboratory study involved a baseline night, a night of sleep loss (approximately 40 h) followed by 5 nighttime recovery sleep periods. Recovery consisted of either five 9-h sleep opportunities or five 6-h sleep opportunities. SETTING: All data collection took place in a controlled laboratory environment at the Centre for Sleep Research. PARTICIPANTS: A total of n = 20 healthy adults (18-35 yrs) participated in the study. RESULTS: Each sleep period was recorded using a standard polysomnographic EEG montage. Waking functions were assessed every 2 hours during all wake periods, using a 10 minute psychomotor vigilance task (PVT) and a subjective alertness visual analogue scale (VAS). Sleep analyses indicated that across the week TST, SOL, REM, and sleep efficiency varied significantly between the 2 conditions, but amounts of SWS did not. Waking functions in the 9-h condition recovered after one sleep period. In the 6-h condition however, mean response time on the PVT was 10% below baseline and subjective alertness 20% below baseline for the entire recovery period. CONCLUSIONS: The results suggest that TST is a key factor in determining recovery. When recovery opportunity is restricted, both sleep variables and measures of waking functions do not recover.     

The forty-eight hour day.

Four normal young adult male subjects were evaluated in a systematically imposed regime of 32 hr of wakefulness and 16 hr of sleep time in an environment free from time cues. Electroencephalographic and electrooculographic recordings were made continuously during the experiment, which lasted for 10 complete cycles. Sleep efficiency was assessed by determining the percentage of sleep time during the assigned sleep period. The average sleep efficiency for the experimental period was 77%. Results, in general, conformed to earlier findings of non-24 hr schedules of sleep and waking: the overall sleep system remains relatively stable across a variety of scheduled variations; however, utilization of the sleep period becomes less efficient as the schedule increasingly deviates from the normal approximately 16 hr wakefulness/8 hr sleep schedule.     

An ultra short episode of sleep is sufficient to promote declarative memory performance

Various studies have demonstrated that a night of sleep has a beneficial effect on the retention of previously acquired declarative material. In two experiments, we addressed the question of whether this effect extends to daytime naps. In the first experiment we assessed free recall of a list of 30 words after a 60 min retention interval that was either filled with daytime napping or waking activity. Memory performance was significantly enhanced after napping as opposed to waking but was not correlated with time spent in slow wave sleep or total sleep time within the napping condition. The second experiment was designed to clarify the role of total sleep time and therefore included an additional third group, which was allowed to nap for no longer than 6 min on average. In comparing word recall after conditions of no napping (waking), short napping, and long napping, we found superior recall for both nap conditions in contrast to waking as well as for long naps in contrast to short naps. These results demonstrate that even an ultra short period of sleep is sufficient to enhance memory processing. We suggest that the mere onset of sleep may initiate active processes of consolidation which - once triggered - remain effective even if sleep is terminated shortly thereafter.     

Effects of the selective 5-HT1B agonist, CGS 12066B, on sleep/waking stages and EEG power spectrum in rats

SUMMARY  Sleep/waking stages and EEG power spectra were studied in rats for 8 h following intraperitoneal administration of CGS 12066B, a selective 5-HT_1B agonist. Waking was increased and rapid eye movement (REM) sleep decreased in a dose-dependent manner. Total slow-wave sleep (TSWS) was reduced, but only in the first 2 h period. The latencies to REM sleep and stable sleep were increased dose-dependently. The drug also induced profound behavioural changes that may account for some of the sleep/waking changes. EEG power densities in waking and TSWS were reduced dose-dependently from 7 to 20 Hz after CGS 12066B, suggesting a tendency towards general deactivation. The increase in waking together with a general deactivation suggest complex effects of CGS 12066B on the sleep/waking axis.     

Consolidation of strictly episodic memories mainly requires rapid eye movement sleep

STUDY OBJECTIVES: The aim of this study is to examine the effects of sleep deprivation during the first or second half of the night on episodic memory consolidation. Episodic memory is defined as memory for events located in time and space. It is also characterized by autonoetic consciousness, which gives a subject the conscious sensation of traveling back in time to relive the original event and forward into the future. DESIGN: Consolidation of episodic information was tested after 4-hour retention intervals, which followed learning and occurred during either the early or late half night, respectively dominated by slow wave sleep (SWS) or rapid eye movement (REM) sleep, or corresponding periods of wakefulness. SETTING: Data collection occurred in the sleep laboratory. PARTICIPANTS: Forty-three young healthy subjects: 9 men and 34 women, age ranging from 18 to 26 years (mean 20.18 +/- 1.94 years) were included in this study. INTERVENTIONS: Waking after a 4-hour retention interval filled with early or late sleep, or 4-hour sleep deprivation, during early or late period of night. MEASUREMENTS AND RESULTS: The cognitive task, named the What-Where-When test, was specially designed to assess factual, spatial, and temporal components of episodic memory. This task was associated with the Remember/Know paradigm to assess autonoetic consciousness. We measured performance on immediate free recall, delayed free recall (after a 4-hour interval of wakefulness or sleep), and delayed recognition. We also calculated a forgetting rate for each feature (factual, spatial, and temporal) and, for the recognition task, scores of autonoetic consciousness (R responses). REM-sleep deprivation was associated with significantly lower recall of spatial information compared to SWS deprivation (P < .01) or late sleep (P < .05) conditions. REM-sleep deprivation was also associated with a higher forgetting rate of temporal information as compared to the early sleep condition (P< .01). Finally, REM-sleep deprivation led subjects to give significantly fewer R responses, indicative of true memories, as compared to SWS deprivation (P < .05). CONCLUSIONS: These results suggest that consolidation of truly episodic memories mainly involves REM sleep.     

The sleeping and waking states of infants: correlations across time and person

Study of the sleeping and waking states of infants was extended to the time domain by analyzing their temporal covariation over a 4-week interval using the intra-person correlation coefficient. As a methodological question, inter-person and intra-person correlation matrices were compared. Twenty-eight fullterm, normal infants were observed in the home weekly from 2 to 5 weeks of age. Over each 7-hour observation, the following states were recorded every 10 seconds: Alert, Non-Alert Waking Activity, Fussing, Crying, Daze, Drowse, Sleep-Wake Transition, Active Sleep, Quiet Sleep, and Unclassified Sleep (sleep during periods of the day when a mother was holding her baby). Data were obtained in two social contexts: when the baby was with the mother and when alone. Significant individual differences were found for each of the states in both social contexts. When the baby was alone, there were a greater number of intra-person correlations than inter-person correlations. All correlations between the waking states and Active Sleep or Quiet Sleep were negative, suggesting the hypothesis that the underlying neurobiological mechanisms may allow a degree of substitution among coupled sleep and wake states. This is an extension of the original hypothesis by Roffwarg, Muzio, and Dement. There were fewer systematic relationships among the states when the babies were with their mother in comparison with periods when they were alone.(ABSTRACT TRUNCATED AT 250 WORDS)     

Facilitated Recall Following RFM And NREM Naps

A novel procedure was used to assess (a) the differential effects of REM and SWS on retention and (b) whether these sleep stages enhanced recall for different types of information. In a 2 × 2 factorial design, subjects were given non-meaningful and meaningful verbal learning tasks. Learning took place either early in the morning or late in the afternoon. Following learning subjects either slept or remained awake for a 2-hr retention interval, after which recall was assessed. Sleep for morning subjects was composed of stages 1, 2 and rapid eye movement sleep while sleep for afternoon subjects was composed of stages 1, 2 and slow wave sleep. Retention for both tasks was better following periods of sleep than following periods of wakefulness but no differences in recall were observed after REM vs. NREM naps.     

Effect of total sleep deprivation on the dimensional complexity of the waking EEG

STUDY OBJECTIVES: Sleep deprivation can affect the waking EEG that may reflect information processing of the brain. We examined the effect of total sleep deprivation (TSD) on nonlinear dynamics of the waking EEG. DESIGN: Paired-group design. SETTING: A sleep disorders laboratory in a hospital. PARTICIPANTS: Twenty healthy male volunteers. INTERVENTIONS: Waking EEG data were recorded from subjects with eyes closed after (a) an 8-hour night's sleep and (b) TSD for 24 hours. The dimensional complexity (D2), as a nonlinear measure of complexity, of the EEG after a full night sleep were compared with those of the EEG after TSD. MEASUREMENTS AND RESULTS: The sleep-deprived states had lower D2 values at three channels (P4, O2, and C3) than normal states. CONCLUSIONS: TSD results in the decrease of complexity in the brain, which may imply sub-optimal information processing of the cerebral cortex. We suggest that the investigation of the relation between nonlinear dynamics of the waking EEG induced by TSD and cognitive performance may offer fruitful clues for understanding the role of sleep and the effects of sleep deprivation on brain function.     

Inception of a 24-hr capacity memory in two mouse strains

Separate groups of C3H and Swiss-Webster (S-W) mice received 25 escape training trials in a straight alley at 8 or 10 days of age. Each of these groups was then divided into 3 retention groups, receiving an additional 25 training trials, 1, 6, or 24 hr following original training. Comparisons of the retest scores with original training scores, as well as with litter-mate controls without prior training, indicated that both strains trained at 8 days of age failed to demonstrate retention of learning when retested 24 hr after training, although they showed reliable 1-hr retention and some evidence of 6-hr retention. However, mice trained at 10 days of age performed better on the retention tests at all retest intervals than they had on original training, as well as better than maturation controls without prior training. These results indicate that a 24-hr retention capacity for escape training develops between 8-10 days of age in both mouse strains.     

Baseline sleep-wake patterns in the pointer dog.

The 24-hr electrographic patterns (EEG, EOG, EMG) of six normal pointer dogs were recorded in a laboratory setting. Two states of sleep (slow-wave and rapid eye movement) and wakefulness (alert and drowsy) were identified. The total recording period comprised 44% of alert wakefulness, 21% of the drowsy state, while slow-wave sleep occupied 23% and REM sleep 12% of the time. The mean length of a REM sleep episode averaged 6 min and the mean REM sleep cycle was 20 min. The mean polycyclic sleep-wake cycle was 83 min. Sleep episodes averaged 45 min and the mean waking episode was 38 min. There was an average of two REM sleep episodes per sleep-wake cycle. The dog has a propensity to sleep over a 16-hr interval from 1300 to 0500 but the most sleep occurred between 2100 and 0400 hr during darkness.     

Reduced Sleep Acutely Influences Sedentary Behavior and Mood But Not Total Energy Intake in Normal-Weight and Obese Women

Using a crossover design, 22 normal-weight and 22 obese women completed two free-living sleep conditions: (a) Normal Sleep: night of ~8 hr time in bed; and (b) Reduced Sleep: night of < 5 hr time in bed). Outcome measures were energy intake, physical activity and sedentary time, and mood. Sleep time was 7.7 ± 0.3 and 4.8 ± 0.2 hrs during the Normal Sleep and Reduced Sleep conditions, respectively (F = 1791.94; p < 0.0001). Energy intake did not differ between groups or as a function of sleep condition (F = 2.46; p = 0.1244). Sedentary time was ~ 30 min higher after the Reduced Sleep condition (F = 4.98; p = 0.0318); other physical activity outcomes were not different by condition (p > 0.05). Total mood score, depression, anger, vigor, fatigue, and confusion were worse after Reduced Sleep (p < 0.05). Reducing sleep acutely and negatively influenced sedentary time and mood in normal-weight and obese women.     

Sleep and rest facilitate auditory learning

Sleep is superior to waking for promoting performance improvements between sessions of visual perceptual and motor learning tasks. Few studies have investigated possible effects of sleep on auditory learning. A key issue is whether sleep specifically promotes learning, or whether restful waking yields similar benefits. According to the "interference hypothesis," sleep facilitates learning because it prevents interference from ongoing sensory input, learning and other cognitive activities that normally occur during waking. We tested this hypothesis by comparing effects of sleep, busy waking (watching a film) and restful waking (lying in the dark) on auditory tone sequence learning. Consistent with recent findings for human language learning, we found that compared with busy waking, sleep between sessions of auditory tone sequence learning enhanced performance improvements. Restful waking provided similar benefits, as predicted based on the interference hypothesis. These findings indicate that physiological, behavioral and environmental conditions that accompany restful waking are sufficient to facilitate learning and may contribute to the facilitation of learning that occurs during sleep.     

The effects of total sleep deprivation on brain functional organization: mutual information analysis of waking human EEG.

Sleep deprivation can affect the waking electroencephalogram (EEG) that may reflect functional organization of the brain. We examined the effect of total sleep deprivation (TSD) on functional organization between different cortical areas from the waking EEG. Waking EEG data were recorded from 18 healthy male volunteers with eyes closed after 8-h night's sleep and after 24 h of TSD. The averaged cross mutual information (A-CMI) after 24 h of TSD were compared to before TSD. 24 h of TSD yielded the decreased A-CMIs in the inter-hemispheric C3-F4, C3-F8, and C3-C4 pairs: therefore, the electrodes that contribute to pairs with significant decrease of A-CMI were C3, F4, F8, and C4. The decreased A-CMIs between C3 and right frontal and central brain areas after 24 h of TSD may reflect the changes of cortico-cortical functional organization by homeostatic process during TSD. Our results of the frontal-area-related A-CMI decreases may support that the frontal brain regions are related to the homeostatic deterioration of brain function due to TSD.     

Effects of early and late nocturnal sleep on priming and spatial memory.

A wordstem priming task (nondeclarative memory), and a mental spatial rotation task (declarative memory) were presented to subjects of an experimental "sleep" group (n = 11) and of a "wake" control group (n = 10). Repetition priming effects and recall of spatial memory were tested after 3-hr retention intervals, which followed learning and were placed either in the early or in the late half of the night. Sleep group subjects slept during the retention intervals while subjects of the wake group stayed awake. As expected, early retention sleep was dominated by slow wave sleep (SWS), whereas rapid eye movement (REM) sleep prevailed during late retention sleep. After early retention sleep, recall of spatial memory was superior to that after late retention sleep (p < 0.01), and also to that after retention intervals of wakefulness (p < 0.05). In contrast, priming was more effective after late than early retention sleep (p < 0.05). It appears that early sleep dominated by SWS facilitates consolidation of declarative memory whereas late sleep dominated by REM sleep facilitates consolidation of nondeclarative memory.     

The effects of handling in infancy on behavioral states in the rabbit

Experimental rabbits were handled for the first 20 days of life while controls were not disturbed. At 25 and 30 days of age, each animal was observed alone for one hr and behavioral states were recorded. They were given the open-field test on Days 31–34. Handling significantly affected the organization of behavioral states and open-field performance. At 25 days handled animals spent a larger percent of sleep time in Active Sleep, and they had a greater percent of REMs than did controls. Experimental rabbits had more Active Waking and less Quiet Waking than controls at 30 days; the mean duration of time spent in Active Waking was greater than that of controls at 30 days, while the mean duration of time spent in Quiet Waking was less than controls.     

Electroencephalogram power density and slow wave sleep as a function of prior waking and circadian phase

Human sleep electroencephalograms, recorded in four experiments, were subjected to spectral analysis. Waking prior to sleep varied from 12 to 36 h and sleep was initiated at different circadian phases. Power density of delta and theta frequencies in rapid-eye-movement (REM) sleep and non-REM (NREM) sleep increased monotonically as a function of prior waking. The increase of power density in the theta frequencies contrasts with the reported decrease of theta activity as detected by period-amplitude analysis. Slow wave activity (power density, 0.25-4.0 Hz) in NREM sleep during the first 3 h of sleep did not deviate significantly from the homeostatic process S of the two-process model of sleep regulation. In contrast, visually scored slow wave sleep, stages 3 and 4, deviated from this prediction at some circadian phases. It is concluded that, in accordance with the two-process model of sleep regulation, slow wave activity in NREM sleep depends on prior waking and is not significantly influenced by circadian phase.