Are we really unconscious in "unconscious" states? Common assumptions revisited.

In the field of consciousness science, there is a tradition to categorize certain states such as slow-wave non-REM sleep and deep general anesthesia as "unconscious". While this categorization seems reasonable at first glance, careful investigations have revealed that it is not so simple. Given that (1) behavioral signs of (un-)consciousness can be unreliable, (2) subjective reports of (un-)consciousness can be unreliable, and, (3) states presumed to be unconscious are not always devoid of reported experience, there are reasons to reexamine our traditional assumptions about "states of unconsciousness". While these issues are not novel, and may be partly semantic, they have implications both for scientific progress and clinical practice. We suggest that focusing on approaches that provide a more pragmatic and nuanced characterization of different experimental conditions may promote clarity in the field going forward, and help us build stronger foundations for future studies.

Random Practice Enhances Retention and Spatial Transfer in Force Field Adaptation.

The contextual-interference effect is a frequently examined phenomenon in motor skill learning but has not been extensively investigated in motor adaptation. Here, we first tested experimentally if the contextual-interference effect is detectable in force field adaptation regarding retention and spatial transfer, and then fitted state-space models to the data to relate the findings to the "forgetting-and-reconstruction hypothesis". Thirty-two participants were divided into two groups with either a random or a blocked practice schedule. They practiced reaching to four targets and were tested 10 min and 24 h afterward for motor retention and spatial transfer on an interpolation and an extrapolation target, and on targets which were shifted 10 cm away. The adaptation progress was participant-specifically fitted with 4-slow-1-fast state-space models accounting for generalization and set breaks. The blocked group adapted faster (p = 0.007) but did not reach a better adaptation at practice end. We found better retention (10 min), interpolation transfer (10 min), and transfer to shifted targets (10 min and 24 h) for the random group (each p < 0.05). However, no differences were found for retention or for the interpolation target after 24 h. Neither group showed transfer to the extrapolation target. The extended state-space model could replicate the behavioral results with some exceptions. The study shows that the contextual-interference effect is partially detectable in practice, short-term retention, and spatial transfer in force field adaptation; and that state-space models provide explanatory descriptions for the contextual-interference effect in force field adaptation.

EEG Lempel-Ziv complexity varies with sleep stage, but does not seem to track dream experience.

In a recent electroencephalography (EEG) sleep study inspired by complexity theories of consciousness, we found that multi-channel signal diversity progressively decreased from wakefulness to slow wave sleep, but failed to find any significant difference between dreaming and non-dreaming awakenings within the same sleep stage (NREM2). However, we did find that multi-channel Lempel-Ziv complexity (LZC) measured over the posterior cortex increased with more perceptual ratings of NREM2 dream experience along a thought-perceptual axis. In this follow-up study, we re-tested our previous findings, using a slightly different approach. Partial sleep-deprivation was followed by evening sleep experiments, with repeated awakenings and immediate dream reports. Participants reported whether they had been dreaming, and were asked to rate how diverse, vivid, perceptual, and thought-like the contents of their dreams were. High density (64 channel) EEG was recorded throughout the experiment, and mean single-channel LZC was calculated for each 30 s sleep epoch. LZC progressively decreased with depth of non-REM sleep. Surprisingly, estimated marginal mean LZC was slightly higher for NREM1 than for wakefulness, but the difference did not remain significant after adjusting for multiple comparisons. We found no significant difference in LZC between dream and non-dream awakenings, nor any significant relationship between LZC and subjective ratings of dream experience, within the same sleep stage (NREM2). The failure to reproduce our own previous finding of a positive correlation between posterior LZC and more perceptual dream experiences, or to find any other correlation between brain signal complexity and subjective experience within NREM2 sleep, raises the question of whether EEG LZC is really a reliable correlate of richness of experience as such, within the same sleep stage.

EEG Signal Diversity Varies With Sleep Stage and Aspects of Dream Experience.

Several theories link consciousness to complex cortical dynamics, as suggested by comparison of brain signal diversity between conscious states and states where consciousness is lost or reduced. In particular, Lempel-Ziv complexity, amplitude coalition entropy and synchrony coalition entropy distinguish wakefulness and REM sleep from deep sleep and anesthesia, and are elevated in psychedelic states, reported to increase the range and vividness of conscious contents. Some studies have even found correlations between complexity measures and facets of self-reported experience. As suggested by integrated information theory and the entropic brain hypothesis, measures of differentiation and signal diversity may therefore be measurable correlates of consciousness and phenomenological richness. Inspired by these ideas, we tested three hypotheses about EEG signal diversity related to sleep and dreaming. First, diversity should decrease with successively deeper stages of non-REM sleep. Second, signal diversity within the same sleep stage should be higher for periods of dreaming vs. non-dreaming. Third, specific aspects of dream contents should correlate with signal diversity in corresponding cortical regions. We employed a repeated awakening paradigm in sleep deprived healthy volunteers, with immediate dream report and rating of dream content along a thought-perceptual axis, from exclusively thought-like to exclusively perceptual. Generalized linear mixed models were used to assess how signal diversity varied with sleep stage, dreaming and thought-perceptual rating. Signal diversity decreased with sleep depth, but was not significantly different between dreaming and non-dreaming, even though there was a significant positive correlation between Lempel-Ziv complexity of EEG recorded over the posterior cortex and thought-perceptual ratings of dream contents.

Contextual Interference Effect Is Independent of Retroactive Inhibition but Variable Practice Is Not Always Beneficial.

Positive effects of variable practice conditions on subsequent motor memory consolidation and generalization are widely accepted and described as the contextual interference effect (CIE). However, the general benefits of CIE are low and these benefits might even depend on decreased retest performances in the blocked-practicing control group, caused by retroactive inhibition. The aim of this study was to investigate if CIE represents a true learning phenomenon or possibly reflects confounding effects of retroactive inhibition. We tested 48 healthy human participants adapting their reaching movements to three different force field magnitudes. Subjects practiced the force fields in either a Blocked (B), Random (R), or Constant (C) schedule. In addition, subjects of the Blocked group performed either a retest schedule that did (Blocked-Matched; BM) or did not (Blocked-Unmatched; BU) control for retroactive inhibition. Results showed that retroactive inhibition did not affect the results of the BU group much and that the Random group showed a better consolidation performance compared to both Blocked groups. However, compared to the Constant group, the Random group showed only slight benefits in its memory consolidation of the mean performance across all force field magnitudes and no benefits in absolute performance values. This indicates that CIE reflects a true motor learning phenomenon, which is independent of retroactive inhibition. However, random practice is not always beneficial over constant practice.

Variable training but not sleep improves consolidation of motor adaptation.

How motor memory consolidates still remains elusive. Consolidation of motor skills has been shown to depend on periods of sleep. Conversely, motor adaptation during tasks not dependent on the hippocampus may not depend on sleep. Some research suggests that the training schedule affects the sleep dependency of motor adaptation tasks. Here, we investigated whether sleep differentially affects memory consolidation that depends on the training schedule. Healthy men were trained with their dominant, right hand on a force-field adaptation task and re-tested after an 11-h consolidation period involving overnight sleep (Sleep) or daytime wakefulness (Wake). Retesting included a transfer test of the non-dominant hand. Half of the subjects in each group adapted to different force-field magnitudes during training with low inter-trial force variability (Sleep-Blocked; Wake-Blocked), and the other half were trained with a high-variability schedule (Sleep-Random; Wake-Random). EEG was recorded during task execution and overnight polysomnography. Consolidation was comparable between Wake and Sleep groups, although performance changes over sleep correlated with sleep spindles nesting in slow-wave upstates. Higher training variability improved retest performance, including transfer learning, and these improvements correlated with higher alpha power in contralateral parietal areas. These enhanced consolidation effects might be fostered by feedback rather than feedforward mechanisms.

Mechanisms within the Parietal Cortex Correlate with the Benefits of Random Practice in Motor Adaptation.

The motor learning literature shows an increased retest or transfer performance after practicing under unstable (random) conditions. This random practice effect (also known as contextual interference effect) is frequently investigated on the behavioral level and discussed in the context of mechanisms of the dorsolateral prefrontal cortex and increased cognitive efforts during movement planning. However, there is a lack of studies examining the random practice effect in motor adaptation tasks and, in general, the underlying neural processes of the random practice effect are not fully understood. We tested 24 right-handed human subjects performing a reaching task using a robotic manipulandum. Subjects learned to adapt either to a blocked or a random schedule of different force field perturbations while subjects' electroencephalography (EEG) was recorded. The behavioral results showed a distinct random practice effect in terms of a more stabilized retest performance of the random compared to the blocked practicing group. Further analyses showed that this effect correlates with changes in the alpha band power in electrodes over parietal areas. We conclude that the random practice effect in this study is facilitated by mechanisms within the parietal cortex during movement execution which might reflect online feedback mechanisms.

Consecutive learning of opposing unimanual motor tasks using the right arm followed by the left arm causes intermanual interference.

Intermanual transfer (motor memory generalization across arms) and motor memory interference (impairment of retest performance in consecutive motor learning) are well-investigated motor learning phenomena. However, the interplay of these phenomena remains elusive, i.e., whether intermanual interference occurs when two unimanual tasks are consecutively learned using different arms. Here, we examine intermanual interference when subjects consecutively adapt their right and left arm movements to novel dynamics. We considered two force field tasks A and B which were of the same structure but mirrored orientation (B = -A). The first test group (ABA-group) consecutively learned task A using their right arm and task B using their left arm before being retested for task A with their right arm. Another test group (AAA-group) learned only task A in the same right-left-right arm schedule. Control subjects learned task A using their right arm without intermediate left arm learning. All groups were able to adapt their right arm movements to force field A and both test groups showed significant intermanual transfer of this initial learning to the contralateral left arm of 21.9% (ABA-group) and 27.6% (AAA-group). Consecutively, both test groups adapted their left arm movements to force field B (ABA-group) or force field A (AAA-group). For the ABA-group, left arm learning caused significant intermanual interference of the initially learned right arm task (68.3% performance decrease). The performance decrease of the AAA-group (10.2%) did not differ from controls (15.5%). These findings suggest that motor control and learning of right and left arm movements involve partly similar neural networks or underlie a vital interhemispheric connectivity. Moreover, our results suggest a preferred internal task representation in extrinsic Cartesian-based coordinates rather than in intrinsic joint-based coordinates because interference was absent when learning was performed in extrinsically equivalent fashion (AAA-group) but interference occurred when learning was performed in intrinsically equivalent fashion (ABA-group).

Bilateral practice improves dominant leg performance in long jump.

Benefits of bilateral practice both for the non-dominant and for the dominant body side have been shown in several studies. Thereby, most of the studies included movement tasks of the upper extremity or investigated sports games in which the ability of acting bilaterally is an essential basis for success and, thus, a bilateral practice is reasonable anyway. Individual unilaterally performed sports including movement tasks of the lower extremity are rarely investigated. Therefore, the aim of our study was to test if contralateral transfer due to bilateral practice can be found in an unilaterally performed sport including the lower extremity. We trained and tested 61 adolescent athletes in long jump to compare the jumping performance of the dominant leg after a 12-week practice period between two groups: a bilateral practice group that practiced specific long jump exercises with both the dominant and non-dominant leg and an unilateral practice group that practiced specific long jump exercises only with the dominant leg. Results showed a superior effect of bilateral practice compared to unilateral practice regarding the jumping performance of the dominant leg. The performance increase at post-test and retention-test for the dominant limb was significantly higher for the bilateral practice group (pre-to-post: 5.2%, pre-to-retention: 7.4%) compared to the unilateral practice group (pre-to-post: 3.4%, pre-to-retention: 4.5%). Thus, bilateral practice should be established in the early practice programmes of track and field athletes to improve the performance of the dominant take-off leg.

Increased gamma band power during movement planning coincides with motor memory retrieval.

The retrieval of motor memory requires a previous memory encoding and subsequent consolidation of the specific motor memory. Previous work showed that motor memory seems to rely on different memory components (e.g., implicit, explicit). However, it is still unknown if explicit components contribute to the retrieval of motor memories formed by dynamic adaptation tasks and which neural correlates are linked to memory retrieval. We investigated the lower and higher gamma bands of subjects' electroencephalography during encoding and retrieval of a dynamic adaptation task. A total of 24 subjects were randomly assigned to a treatment and control group. Both groups adapted to a force field A on day 1 and were re-exposed to the same force field A on day 3 of the experiment. On day 2, treatment group learned an interfering force field B whereas control group had a day rest. Kinematic analyses showed that control group improved their initial motor performance from day 1 to day 3 but treatment group did not. This behavioral result coincided with an increased higher gamma band power in the electrodes over prefrontal areas on the initial trials of day 3 for control but not treatment group. Intriguingly, this effect vanished with the subsequent re-adaptation on day 3. We suggest that improved re-test performance in a dynamic motor adaptation task is contributed by explicit memory and that gamma bands in the electrodes over the prefrontal cortex are linked to these explicit components. Furthermore, we suggest that the contribution of explicit memory vanishes with the subsequent re-adaptation while task automaticity increases.

Intermanual transfer characteristics of dynamic learning: direction, coordinate frame, and consolidation of interlimb generalization.

Intermanual transfer, i.e., generalization of motor learning across hands, is a well-accepted phenomenon of motor learning. Yet, there are open questions regarding the characteristics of this transfer, particularly the intermanual transfer of dynamic learning. In this study, we investigated intermanual transfer in a force field adaptation task concerning the direction and the coordinate frame of transfer as well as the influence of a 24-h consolidation period on the transfer. We tested 48 healthy human subjects for transfer from dominant to nondominant hand, and vice versa. We considered two features of transfer. First, we examined transfer to the untrained hand using force channel trials that suppress error feedback and learning mechanisms to assess intermanual transfer in the form of a practice-dependent bias. Second, we considered transfer by exposing the subjects to the force field with the untrained hand to check for faster learning of the dynamics (interlimb savings). Half of the subjects were tested for transfer immediately after adaptation, whereas the other half were tested after a 24-h consolidation period. Our results showed intermanual transfer both from dominant to nondominant hand and vice versa in extrinsic coordinates. After the consolidation period, transfer effects were weakened. Moreover, the transfer effects were negligible compared with the subjects' ability to rapidly adapt to the force field condition. We conclude that intermanual transfer is a bidirectional phenomenon that vanishes with time. However, the ability to transfer motor learning seems to play a minor role compared with the rapid adaptation processes.