Effects of acute bouts of evening resistance or endurance exercises on sleep EEG and salivary cortisol.

Introduction: Deleterious effects of exercise close to bedtime could be due to increased physiological arousal that can be detected during sleep using sleep spectral analysis. Resistance and endurance exercises have different effects on cortisol release that may lead them to impact sleep spectral signatures differently. The present study aimed to investigate the effects of two types of evening exercise on sleep architecture, sleep spectral parameters and salivary cortisol. Methods: Young healthy participants came to our laboratory to undergo 3 counterbalanced pre-sleep conditions that started 1 h before bedtime (a resistance and an endurance exercise conditions of 30 min duration, identical in terms of workload; and a control condition) followed by polysomnographic recordings. Results were compared between the three conditions for 16 participants. Results: Sleep efficiency was lower after both endurance and resistance exercise than after the control condition. Total sleep time was lower after endurance exercise compared to the control condition. Sleep spectral analyses showed that both endurance and resistance exercises led to greater alpha power during N1 sleep stage and greater theta power during N2 sleep stage compared to the control condition. The endurance exercise led to greater beta power during N2 sleep stage, greater alpha power during REM sleep, and higher cortisol levels compared to the control condition (trend), and compared to the resistance exercise condition (significant). The resistance exercise led to lower beta power during N2 sleep stage than the control condition and lower cortisol levels than the endurance exercise condition. Discussion: This study underlines significant modifications of sleep quality and quantity after both moderate evening endurance and resistance exercises. Still, these effects cannot be considered as deleterious. In contrast to the resistance exercise, endurance exercise led to an increase in sleep EEG activity associated with hyperarousal during sleep and higher cortisol levels, suggesting an hyperarousal effect of endurance exercise performed in the evening. These results align with previous warning about the arousal effects of evening exercise but do not support the notion of deleterious effects on sleep. While these results provide support for the physiological effects of evening exercises on sleep, replication with larger sample size is needed.

Effects of sleep disturbances and circadian rhythms modifications on cognition in breast cancer women before and after adjuvant chemotherapy: the ICANSLEEP-1 protocol.

BACKGROUND: Many patients treated for breast cancer (BC) complain about cognitive difficulties affecting their daily lives. Recently, sleep disturbances and circadian rhythm disruptions have been brought to the fore as potential contributors to cognitive difficulties in patients with BC. Yet, studies on these factors as well as their neural correlates are scarce. The purpose of the ICANSLEEP-1 (Impact of SLEEP disturbances in CANcer) study is to characterize sleep using polysomnography and its relationship with the evolution of cognitive functioning at both the behavioral and the neuroanatomical levels across treatment in BC patients treated or not with adjuvant chemotherapy. METHODS: ICANSLEEP-1 is a longitudinal study including BC patients treated with adjuvant chemotherapy (n = 25) or not treated with adjuvant chemotherapy (n = 25) and healthy controls with no history of BC (n = 25) matched for age (45-65 years old) and education level. The evaluations will take place within 6 weeks after inclusion, before the initiation of chemotherapy (for BC patients who are candidates for chemotherapy) or before the first fraction of radiotherapy (for BC patients with no indication for chemotherapy) and 6 months later (corresponding to 2 weeks after the end of chemotherapy). Episodic memory, executive functions, psychological factors, and quality of life will be assessed with validated neuropsychological tests and self-questionnaires. Sleep quantity and quality will be assessed with polysomnography and circadian rhythms with both actigraphy and saliva cortisol. Grey and white matter volumes, as well as white matter microstructural integrity, will be compared across time between patients and controls and will serve to further investigate the relationship between sleep disturbances and cognitive decline. DISCUSSION: Our results will help patients and clinicians to better understand sleep disturbances in BC and their relationship with cognitive functioning across treatment. This will aid the identification of more appropriate sleep therapeutic approaches adapted to BC patients. Improving sleep in BC would eventually help limit cognitive deficits and thus improve quality of life during and after treatments. TRIAL REGISTRATION: NCT05414357, registered June 10, 2022. PROTOCOL VERSION: Version 1.2 dated March 23, 2022.

Intact memory storage but impaired retrieval in visual memory in autism: New insights from an electrophysiological study.

In a recent study on visual episodic memory (Desaunay, Clochon, et al., 2020), we have shown event-related potentials (ERPs) differences associated with priming (150-300 msec), familiarity (350-470 msec), and recollection (600-700 msec), in young people with autism spectrum disorders (ASD) compared with typical development (TD). To go further into the study of the processes of storage and retrieval of the memory trace, we re-analyzed Desaunay, Clochon, et al's data using time-frequency analysis, that is, event-related synchronization and desynchronization (ERS/ERD). This allows a decomposition of the spectral power within frequency bands associated with these ERPs. We focused both on the same time windows and the same regions of interest as previously published. We mainly identified, in ASD compared with TD, reduced ERS in low-frequencies (delta, theta) in early time-windows, and non-significant differences in ERD in higher frequencies (alpha, beta1) in all time-windows. Reduced ERS during recognition confirmed previously reported diminution of priming effects and difficulties in manipulation and retrieval of both semantic and episodic information. Conversely, preserved ERD corroborates a preservation of memory storage processes. These observations are consistent with a cognitive model of memory in ASD, that suggests difficulties in cognitive operations or executive demand at retrieval, subsequent to successful long-term storage of information. LAY SUMMARY: We assessed the EEG synchronization and desynchronization, during visual episodic recognition. We observed, in youth with Autism, reduced synchronization in low-frequencies (delta, theta), suggesting reduced access to and manipulation of long-term stored information. By contrast, non-significant differences in desynchronization at higher frequencies (alpha, beta frequency bands), that support long-term stored semantic and episodic information, suggested preserved memory traces.

Attentional capture mediates the emergence and suppression of intrusive memories.

Intrusive memories hijack consciousness and their control may lead to forgetting. However, the contribution of reflexive attention to qualifying a memory signal as interfering is unknown. We used machine learning to decode the brain's electrical activity and pinpoint the otherwise hidden emergence of intrusive memories reported during a memory suppression task. Importantly, the algorithm was trained on an independent attentional model of visual activity, mimicking either the abrupt and interfering appearance of visual scenes into conscious awareness or their deliberate exploration. Intrusion of memories into conscious awareness were decoded above chance. The decoding accuracy increased when the algorithm was trained using a model of reflexive attention. Conscious detection of intrusive activity decoded from the brain signal was central to the future silencing of suppressed memories and later forgetting. Unwanted memories require the reflexive orienting of attention and access to consciousness to be suppressed effectively by inhibitory control.

EEG resting-state functional connectivity: evidence for an imbalance of external/internal information integration in autism.

BACKGROUND: Autism spectrum disorder (ASD) is associated with atypical neural activity in resting state. Most of the studies have focused on abnormalities in alpha frequency as a marker of ASD dysfunctions. However, few have explored alpha synchronization within a specific interest in resting-state networks, namely the default mode network (DMN), the sensorimotor network (SMN), and the dorsal attention network (DAN). These functional connectivity analyses provide relevant insight into the neurophysiological correlates of multimodal integration in ASD. METHODS: Using high temporal resolution EEG, the present study investigates the functional connectivity in the alpha band within and between the DMN, SMN, and the DAN. We examined eyes-closed EEG alpha lagged phase synchronization, using standardized low-resolution brain electromagnetic tomography (sLORETA) in 29 participants with ASD and 38 developing (TD) controls (age, sex, and IQ matched). RESULTS: We observed reduced functional connectivity in the ASD group relative to TD controls, within and between the DMN, the SMN, and the DAN. We identified three hubs of dysconnectivity in ASD: the posterior cingulate cortex, the precuneus, and the medial frontal gyrus. These three regions also presented decreased current source density in the alpha band. CONCLUSION: These results shed light on possible multimodal integration impairments affecting the communication between bottom-up and top-down information. The observed hypoconnectivity between the DMN, SMN, and DAN could also be related to difficulties in switching between externally oriented attention and internally oriented thoughts.

Local Processing Bias Impacts Implicit and Explicit Memory in Autism.

Autism spectrum disorder (ASD) is characterized by atypical perception, including processing that is biased toward local details rather than global configurations. This bias may impact on memory. The present study examined the effect of this perception on both implicit (Experiment 1) and explicit (Experiment 2) memory in conditions that promote either local or global processing. The first experiment consisted of an object identification priming task using two distinct encoding conditions: one favoring local processing (Local condition) and the other favoring global processing (Global condition) of drawings. The second experiment focused on episodic (explicit) memory with two different cartoon recognition tasks that favored either local (i.e., processing specific details) or a global processing (i.e., processing each cartoon as a whole). In addition, all the participants underwent a general clinical cognitive assessment aimed at documenting their cognitive profile and enabling correlational analyses with experimental memory tasks. Seventeen participants with ASD and 17 typically developing (TD) controls aged from 10 to 16 years participated to the first experiment and 13 ASD matched with 13 TD participants were included for the second experiment. Experiment 1 confirmed the preservation of priming effects in ASD but, unlike the Comparison group, the ASD group did not increase his performance as controls after a globally oriented processing. Experiment 2 revealed that local processing led to difficulties in discriminating lures from targets in a recognition task when both lures and targets shared common details. The correlation analysis revealed that these difficulties were associated with processing speed and inhibition. These preliminary results suggest that natural perceptual processes oriented toward local information in ASD may impact upon their implicit memory by preventing globally oriented processing in time-limited conditions and induce confusion between explicit memories that share common details.

Exploring the Event-Related Potentials' Time Course of Associative Recognition in Autism.

Behavioral data on episodic recollection in autism spectrum disorders (ASD) point limited relational memory functioning. However, the involvement of successive memory processes in the profile of episodic memory in ASD needs more study. Here, we used event-related potentials (ERP) to investigate the time course of episodic recollection with an associative recognition paradigm with picture pairs. Twenty-two participants with ASD and 32 with typical development (TD), all right-handed, were included. Behavioral results confirmed difficulties in correctly recognizing identical pairs in the ASD relative to TD group. We found an unexpected amplitude decrement on the P2 (220-270 msec) and FN400 (350-470 msec) potentials, suggesting diminished priming and familiarity effects in the ASD relative to TD group. However, ERP data revealed that the recognition of associative information relies on the same electrophysiological process (old/new effect in the 600-700-msec late positive component) in ASD participants as in TD ones, with a parietal extension in the ASD group. These results suggest that the electrophysiological processes of associative recognition are qualitatively similar in individuals with and without ASD but may differ quantitatively. This difference may be driven by the reduced early processing of picture pairs that may in turn lead to their diminished integration into the semantic memory system, being partially compensated by a greater involvement of associative memory during the recollection process. Other studies would be useful to go further in identifying these cognitive processes involved in atypical recognition in ASD and their neural substrates. Autism Res 2020, 13: 1998-2016. © 2020 International Society for Autism Research and Wiley Periodicals LLC LAY SUMMARY: We identified diminished performance on the associative recognition of picture pairs in adolescents and young adults with autism when compared to typical development. Electrophysiological data revealed qualitative similarities but quantitative differences between-group, with diminished priming and familiarity processes partially compensated by an enhanced parietal recollection process.

Long-term modulation of cardiac activity induced by inhibitory control over emotional memories.

Efforts to exclude past experiences from conscious awareness can lead to forgetting. Memory suppression is central to affective disorders, but we still do not really know whether emotions, including their physiological causes, are also impacted by this process in normal functioning individuals. In two studies, we measured the after-effects of suppressing negative memories on cardiac response in healthy participants. Results of Study 1 revealed that efficient control of memories was associated with long-term inhibition of the cardiac deceleration that is normally induced by disgusting stimuli. Attempts to suppress sad memories, by contrast, aggravated the cardiac response, an effect that was closely related to the inability to forget this specific material. In Study 2, electroencephalography revealed a reduction in power in the theta (3-8 Hz), alpha (8-12 Hz) and low-beta (13-20 Hz) bands during the suppression of unwanted memories, compared with their voluntary recall. Interestingly, however, the reduction of power in the theta frequency band during memory control was related to a subsequent inhibition of the cardiac response. These results provide a neurophysiological basis for the influence of memory control mechanisms on the cardiac system, opening up new avenues and questions for treating intrusive memories using motivated forgetting.

Influence of emotional complexity on the neural substrates of affective theory of mind.

Affective theory of mind (ToM) depends on both the decoding of emotional expressions and the reasoning on emotional mental states from social situations. While previous studies characterized the neural substrates underlying these processes, it remains unclear whether the nature of the emotional state inferred from others can influence the brain activation associated with affective ToM. In the present study, we focused on two types of emotions: basic emotions (BEs) (e.g., anger and surprise), which are innate and universal and self-conscious emotions (e.g., pride and embarrassment), which correspond to a special class of emotions involving the self and including a representation of one's relative reactions to internal and external standards. Specifically, we used an ecological functional MRI paradigm, on 21 healthy young subjects, to compare brain activations during the decoding of and the reasoning on others' self-conscious, basic and neutral mental states. Our results showed that compared to neutral states, the inference of self-conscious and basic emotional states from others elicited more activation in several core regions of affective ToM. Direct comparisons between emotional conditions revealed more activation for self-conscious than BEs in the right temporoparietal junction during the reasoning process and in left middle occipital regions during the decoding process. Further analyses using a localizer task showed that the extrastriate body area was more recruited for decoding others' self-conscious versus BEs, which emphasize the importance of body clues to properly infer these emotions. Using an original task allowing for an ecological assessment of the affective ToM, these results demonstrate that the complexity of the emotion inferred to others can influence the recruitment of ToM network. This study also validates the use of our task as an ecological tool to assess the affective ToM, constituting an avenue for the characterization of ToM impairments in neurological conditions.

Impact of Semantic Relatedness on Associative Memory: An ERP Study.

Encoding and retrieval processes in memory for pairs of pictures are thought to be influenced by inter-item similarity and by features of individual items. Using Event-Related Potentials (ERP), we aimed to identify how these processes impact on both the early mid-frontal FN400 and the Late Positive Component (LPC) potentials during associative retrieval of pictures. Twenty young adults undertook a sham task, using an incidental encoding of semantically related and unrelated pairs of drawings. At test, we conducted a recognition task in which participants were asked to identify target identical pairs of pictures, which could be semantically related or unrelated, among new and rearranged pairs. We observed semantic (related and unrelated pairs) and condition effects (old, rearranged and new pairs) on the early mid-frontal potential. First, a lower amplitude was shown for identical and rearranged semantically related pairs, which might reflect a retrieval process driven by semantic cues. Second, among semantically unrelated pairs, we found a larger negativity for identical pairs, compared to rearranged and new ones, suggesting additional retrieval processing that focuses on associative information. We also observed an LPC old/new effect with a mid-parietal and a right occipito-parietal topography for semantically related and unrelated old pairs, demonstrating a recollection phenomenon irrespective of the degree of association. These findings suggest that associative recognition using visual stimuli begins at early stages of retrieval, and differs according to the degree of semantic relatedness among items. However, either strategy may ultimately lead to recollection processes.

Associative episodic memory and recollective processes in childhood temporal lobe epilepsy.

While the current literature on children suffering from temporal lobe epilepsy (CTLE) mostly focuses on material-related episodic memory deficits according to seizure-onset lateralization, the present study examined associative episodic memory according to the type of information to memorize (e.g., factual, spatial, and sequential) and further investigated subjective and objective recollection. Eleven children with left temporal lobe epilepsy (LTLE), 10 children with right temporal lobe epilepsy (RTLE), among whom 9 displayed hippocampal sclerosis (HS), and 42 healthy controls completed the WHAT-WHEN-WHERE protocol (Guillery-Girard et al., 2013). Group comparisons were first conducted according to the affected side and second according to the underlying pathology. Results showed associative memory impairments in patients irrespective of the affected side. Moreover, this study revealed that HS is particularly deleterious to associative and subjective recollection in CTLE. In addition, this study emphasizes the need for assessing episodic memory in childhood TLE beyond material specificity.

Specific EEG sleep pattern in the prefrontal cortex in primary insomnia.

OBJECTIVE: To assess the specific prefrontal activity in comparison to those in the other main cortical areas in primary insomnia patients and in good sleepers. METHODS: Fourteen primary insomnia patients and 11 good sleepers were included in the analysis. Participants completed one night of polysomnography in the sleep lab. Power spectra were calculated during the NREM (Non-rapid eyes movements) and the REM (Rapid eyes movements) sleep periods at prefrontal, occipital, temporal and central electrode positions. RESULTS: During the NREM sleep, the power spectra did not differ between groups in the prefrontal cortex; while primary insomnia patients exhibited a higher beta power spectrum and a lower delta power spectrum compared to good sleepers in other areas. During the REM sleep, the beta1 power spectrum was lower in the prefrontal cortex in primary insomnia patients compared to good sleepers; while no significant difference between groups was obtained for the other areas. CONCLUSIONS: The present study shows a specific prefrontal sleep pattern during the whole sleep period. In addition, we suggest that primary insomnia patients displayed a dysfunction in the reactivation of the limbic system during the REM sleep and we give additional arguments in favor of a sleep-protection mechanism displayed by primary insomnia patients.

Developmental trajectories of associative memory from childhood to adulthood: a behavioral and neuroimaging study.

Episodic memory refers to the capacity to bind multimodal memories to constitute a unique personal event. Most developmental studies on episodic memory focused on one specific component, i.e., the core factual information. The present study examines the relevance of a novel episodic paradigm to assess its developmental trajectories in a more comprehensive way according to the type of association (item-feature, item-location, and item-sequence associations) with measures of both objective and subjective recollection. We conducted a behavioral study aimed at testing the effects of age in a large sample of 160 children, adolescents, and young adults (6-23 years old). We confronted the behavioral data to the neural correlates in a subgroup of 30 children using voxel-based morphometry. Behavioral data outlined differential developmental trajectories according to the type of association, with a continuous increase of factual associative memory efficiency until 10 years, a linear increase of performance in spatial associative memory that pursues until early adulthood and an abrupt increase in temporal associative memory efficiency between 9 and 10. Regarding recollection, measures showed a more pronounced enhancement from 9 to 10 years. Hence, behavioral data highlight a peculiar period in late childhood (8-10 years old) crucial for the developmental time course of episodic memory. Regarding structural data, we found that the improvement of associative memory efficiency was related to a decrease in gray matter volume in a large cerebral network including the dorsolateral and ventrolateral prefrontal cortex (and superior and anterior temporal regions), and the hippocampus bilaterally. These data suggest that multimodal integration would probably be related to the maturation of temporal regions and modulated by a fronto-parietal network. Besides, our findings emphasize the relevance of the present paradigm to assess episodic memory especially in the clinical setting.

"Disorganized in time": impact of bottom-up and top-down negative emotion generation on memory formation among healthy and traumatized adolescents.

"Travelling in time," a central feature of episodic memory is severely affected among individuals with Post Traumatic Stress Disorder (PTSD) with two opposite effects: vivid traumatic memories are unorganized in temporality (bottom-up processes), non-traumatic personal memories tend to lack spatio-temporal details and false recognitions occur more frequently that in the general population (top-down processes). To test the effect of these two types of processes (i.e. bottom-up and top-down) on emotional memory, we conducted two studies in healthy and traumatized adolescents, a period of life in which vulnerability to emotion is particularly high. Using negative and neutral images selected from the international affective picture system (IAPS), stimuli were divided into perceptual images (emotion generated by perceptual details) and conceptual images (emotion generated by the general meaning of the material). Both categories of stimuli were then used, along with neutral pictures, in a memory task with two phases (encoding and recognition). In both populations, we reported a differential effect of the emotional material on encoding and recognition. Negative perceptual scenes induced an attentional capture effect during encoding and enhanced the recollective distinctiveness. Conversely, the encoding of conceptual scenes was similar to neutral ones, but the conceptual relatedness induced false memories at retrieval. However, among individuals with PTSD, two subgroups of patients were identified. The first subgroup processed the scenes faster than controls, except for the perceptual scenes, and obtained similar performances to controls in the recognition task. The second subgroup group desmonstrated an attentional deficit in the encoding task with no benefit from the distinctiveness associated with negative perceptual scenes on memory performances. These findings provide a new perspective on how negative emotional information may have opposite influences on memory in normal and traumatized individuals. It also gives clues to understand how intrusive memories and overgeneralization takes place in PTSD.

Changes in sleep theta rhythm are related to episodic memory impairment in early Alzheimer's disease.

Impairments have been reported both in sleep structure and episodic memory in Alzheimer's disease [AD]. Our objective was to investigate the relationships between episodic memory deficits and electro-encephalography [EEG] abnormalities occurring during sleep in patients with early AD. Postlearning sleep was recorded in 14 patients with mild to moderate AD, and 14 healthy elderly controls after they performed an episodic memory task derived from the Grober and Buschke's procedure. For each sleep stage, the relative power and mean frequency in each band were analyzed. Relative to agematched controls, AD patients presented faster mean theta frequency in both REM sleep and slow wave sleep [SWS]. In AD patients, a correlative analysis revealed that faster theta frequency during SWS was associated with better delayed episodic recall. We assume that increased theta activity reflects changes in neuronal activity to maintain memory performance, indicating that compensatory mechanisms already described at the waking state could also be engaged during SWS.

A combined neuropsychological and brain imaging study of obstructive sleep apnea.

Patients with obstructive sleep apnea (OSA) show neuropsychological impairments ranging from vigilance decrements, attentional lapses and memory gaps to decreased motor coordination, but their cognitive profile, and the origin of the impairments, remain unclear. We sought to establish the neuropsychological profile of 16 newly diagnosed apneics and to highlight both their morphological and functional brain abnormalities. We used an extensive neuropsychological test battery to investigate attention and vigilance, executive functions, episodic memory and motor domains. For brain imaging, we used the optimized voxel-based morphometry procedure for the MRI data, resting-state (18)F-fluoro-2-deoxy-D-Glucose positron emission tomography ((18)FDG-PET) with correction for partial volume effects (PVEs) and voxel-based analyses. In terms of neurobehavioral performance, our patients displayed objective daytime somnolence but little impairment in memory and motor domains. Cerebral data revealed gray matter loss in the frontal and temporo-parieto-occipital cortices, the thalamus, hippocampal region, some basal ganglia and cerebellar regions, mainly in the right hemisphere. The decrease in brain metabolism was also right-lateralized, but more restricted than the gray matter density changes, and involved the precuneus, the middle and posterior cingulate gyrus, and the parieto-occipital cortex, as well as the prefrontal cortex. To conclude, despite the presence of only minor memory and motor impairments, our patients displayed significant cerebral changes in terms of both gray matter density and metabolic levels, and may have benefited from cognitive reserve and compensatory mechanisms. Thus, cerebral changes in OSA patients may precede the onset of notable neuropsychological consequences.

Structural and metabolic correlates of episodic memory in relation to the depth of encoding in normal aging.

This study set out to establish the relationship between changes in episodic memory retrieval in normal aging on the one hand and gray matter volume and (18)FDG uptake on the other. Structural MRI, resting-state (18)FDG-PET, and an episodic memory task manipulating the depth of encoding and the retention interval were administered to 46 healthy subjects divided into three groups according to their age (young, middle-aged, and elderly adults). Memory decline was found not to be linear in the course of normal aging: Whatever the retention interval, the retrieval of shallowly encoded words was impaired in both the middle-aged and the elderly, whereas the retrieval of deeply encoded words only declined in the elderly. In middle-aged and elderly subjects, the reduced performance in the shallow encoding condition was mainly related to posterior mediotemporal volume and metabolism. By contrast, the impaired retrieval of deeply encoded words in the elderly group was mainly related to frontal and parietal regions, suggesting the adoption of inefficient strategic processes.

Early age-related changes in episodic memory retrieval as revealed by event-related potentials.

Familiarity is better preserved than recollection in ageing. The age at which changes first occur and the slope of the subsequent decline, however, remain unclear. In this study, we investigated changes in episodic memory, by using event-related potentials (ERPs) in young (m=24), middle-aged (m=58) and older (m=70) adults. Although behavioural performance did not change before the age of 65 years, changes in ERP correlates were already present in the middle-aged adults. The ERP correlates of recollection and monitoring processes were the first to be affected by ageing, with a linear decrease as age increased. Conversely, the ERP correlate of familiarity remained unchanged, at least up to the age of 65 years. These results suggest a differential time course for the age effects on episodic retrieval.

Is there a link between sleep changes and memory in Alzheimer's disease?

Aging and Alzheimer's disease (AD) are both characterized by memory impairments and sleep changes. We investigated the potential link between these disturbances, focusing on sleep spindles, involved in memory consolidation. Two episodic memory tasks were given to young and old healthy participants, as well as to AD patients. Postlearning sleep was recorded. Sleep spindles were globally reduced in aging and AD. AD patients also exhibited a further decrease in fast spindles. Besides, mean intensity of fast spindles was positively correlated, in AD patients, with immediate recall performance. Our results are the first report of a specific decrease in fast spindles in AD, associated with learning abilities. They also give further hints for a functional differentiation between slow and fast spindles.