The influence of travel time on perceived traveled distance varies by spatiotemporal scale.

The influence of travel time on perceived traveled distance has often been studied, but the results are inconsistent regarding the relationship between the two magnitudes. We argue that this is due to differences in the lengths of investigated travel distances and hypothesize that the influence of travel time differs for rather short compared to rather long traveled distances. We tested this hypothesis in a virtual environment presented on a desktop as well as through a head-mounted display. Our results show that, for longer distances, more travel time leads to longer perceived distance, while we do not find an influence of travel time on shorter distances. The presentation through an HMD vs. desktop only influenced distance judgments in the short distance condition. These results are in line with the idea that the influence of travel time varies by the length of the traveled distance, and provide insights on the question of how distance perception in path integration studies is affected by travel time, thereby resolving inconsistencies reported in previous studies.

Memory traces of duration and location in the right intraparietal sulcus.

Time and space form an integral part of every human experience, and for the neuronal representation of these perceptual dimensions, previous studies point to the involvement of the right-hemispheric intraparietal sulcus and structures in the medial temporal lobe. Here we used multi-voxel pattern analysis (MVPA) to investigate long-term memory traces for temporal and spatial stimulus features in those areas. Participants were trained on four images associated with short versus long durations and with left versus right locations. Our results demonstrate stable representations of both temporal and spatial information in the right posterior intraparietal sulcus. Building upon previous findings of stable neuronal codes for directly perceived durations and locations, these results show that the reactivation of long-term memory traces for temporal and spatial features can be decoded from neuronal activation patterns in the right parietal cortex.

Rapid calibration to dynamic temporal contexts.

The prediction of future events and the preparation of appropriate behavioural reactions rely on an accurate perception of temporal regularities. In dynamic environments, temporal regularities are subject to slow and sudden changes, and adaptation to these changes is an important requirement for efficient behaviour. Bayesian models have proven a useful tool to understand the processing of temporal regularities in humans; yet an open question pertains to the degree of flexibility of the prior that is required for optimal modelling of behaviour. Here we directly compare dynamic models (with continuously changing prior expectations) and static models (a stable prior for each experimental session) with their ability to describe regression effects in interval timing. Our results show that dynamic Bayesian models are superior when describing the responses to slow, continuous environmental changes, whereas static models are more suitable to describe responses to sudden changes. In time perception research, these results will be informative for the choice of adequate computational models and enhance our understanding of the neuronal computations underlying human timing behaviour.

Validating virtual reality for time perception research: Virtual reality changes expectations about the duration of physical processes, but not the sense of time.

Immersive virtual reality (VR) provides a versatile method for investigating human time perception, because it allows the manipulation and control of relevant variables (e.g., the speed of environmental changes) that cannot be modified in the real world. However, an important premise for interpreting the results of VR studies, namely that the method itself does not affect time perception, has received little attention. Here we tested this assumption by comparing timing performance in a real environment and a VR scenario. Participants performed two timing tasks, requiring the production of intervals defined either by numerical values ("eight seconds") or by a physical process ("the time it takes for a bottle to run out when turned over"). We found that the experience of immersive VR exclusively altered judgments about the duration of physical processes, whereas judgments about the duration of abstract time units were unaffected. These results demonstrate that effects of VR on timing performance are not driven by changes in time perception itself, but rather by altered expectations regarding the duration of physical processes. The present study validates the use of VR in time perception research and strengthens the interpretation of changed timing behaviour induced by manipulations within VR.

Spatial navigation under threat: aversive apprehensions improve route retracing in higher versus lower trait anxious individuals.

Spatial navigation is a basic function for survival, and the ability to retrace a route has direct relevance for avoiding dangerous places. This study investigates the effects of aversive apprehensions on spatial navigation in a virtual urban environment. Healthy participants with varying degrees of trait anxiety performed a route-repetition and a route-retracing task under threatening and safe context conditions. Results reveal an interaction between the effect of threatening/safe environments and trait anxiety: while threat impairs route-retracing in lower-anxious individuals, this navigational skill is boosted in higher-anxious individuals. According to attentional control theory, this finding can be explained by an attentional shift toward information relevant for intuitive coping strategies (i.e., running away), which should be more pronounced in higher-anxious individuals. On a broader scale, our results demonstrate an often-neglected advantage of trait anxiety, namely that it promotes the processing of environmental information relevant for coping strategies and thus prepares the organism for adequate flight responses.

Cross-dimensional interference between time and distance during spatial navigation is mediated by speed representations in intraparietal sulcus and area hMT.

When navigating a straight path, perceived travel time and perceived traveled distance are linked via movement speed. Behavioral studies have revealed systematic interferences between the perception of travel time and distance, but the role of neuronal representations of movement speed for these effects has not been addressed to date. Using a combined fMRI-behavioral paradigm, we investigate the neuronal representations that underlie cross-dimensional interferences between travel time and traveled distance. Participants underwent fMRI while experiencing visual forward movements for either a short or a long duration, and covering either a short or a long distance. At the behavioral level, we found bi-directional interference effects between time and distance perception, which was correlated with greater representational similarity in speed-sensitive brain regions. The strength of the distance-on-time effect scaled with representational similarity in the left human middle temporal complex (hMT+), and the strength of the time-on-distance effect scaled with representational similarity in the right intraparietal sulcus (IPS). In accordance with the idea that the interference is mediated by the perception of speed, distance-on-time and time-on-distance effects were of opposing directions. Increases in traveled distance led to increases in perceived travel time, while increases in travel time led to decreases in perceived traveled distance. Together, these findings support the view that cross-dimensional interference effects between travel time and traveled distance are mediated by neuronal representations of movement speed.

Reducing the tendency for chronometric counting in duration discrimination tasks.

Chronometric counting is a prevalent issue in the study of human time perception as it reduces the construct validity of tasks and can conceal existing timing deficits. Several methods have been proposed to prevent counting strategies, but the factors promoting those strategies in specific tasks are largely uninvestigated. Here, we modified a classical two-interval duration discrimination task in two aspects that could affect the tendency to apply counting strategies. We removed the pause between the two intervals and changed the task instructions: Participants decided whether a short event occurred in the first or in the second half of a reference duration. In Experiment 1, both classical and modified task versions were performed under timing conditions, in which participants were asked not to count, and counting conditions, in which counting was explicitly instructed. The task modifications led to (i) a general decrease in judgment precision, (ii) a shift of the point of subjective equality, and (iii) a counting-related increase in reaction times, suggesting enhanced cognitive effort of counting during the modified task version. Precision in the two task versions was not differently affected by instructed counting. Experiment 2 demonstrates that-in the absence of any counting-related instructions-participants are less likely to engage in spontaneous counting in the modified task version. These results enhance our understanding of the two-interval duration discrimination task and demonstrate that the modifications tested here-although they do not significantly reduce the effectiveness of instructed counting-can diminish the spontaneous tendency to adopt counting strategies.

Temporal context effects are associated with cognitive status in advanced age.

The perception of temporal intervals changes during the life-span, and especially older adults demonstrate specific impairments of timing abilities. Recently, we demonstrated that timing performance and cognitive status are correlated in older adults, suggesting that timing tasks can serve as a behavioral marker for the development of dementia. Easy-to-administer and retest-capable timing tasks therefore have potential as diagnostic tools for tracking cognitive decline. However, before being tested in a clinical cohort study, a further validation and specification of the original findings is warranted. Here we introduce several modifications of the original task and investigated the effects of temporal context on time perception in older adults (> 65 years) with low versus high scores in the Montreal Cognitive Assessment survey (MoCA) and a test of memory functioning. In line with our previous work, we found that temporal context effects were more pronounced with increasing memory deficits, but also that these effects are stronger for realistic compared to abstract visual stimuli. Furthermore, we show that two distinct temporal contexts influence timing behavior in separate experimental blocks, as well as in a mixed block in which both contexts are presented together. These results replicate and extend our previous findings. They demonstrate the stability of the effect for different stimulus material and show that timing tasks can reveal valuable information about the cognitive status of older adults. In the future, these findings could serve as a basis for the development of a diagnostic tool for pathological cognitive decline at an early, pre-clinical stage.

Synthesis of Arylidene-ß-lactams via exo-Selective Matsuda-Heck Arylation of Methylene-ß-lactams.

exo-Methylene-ß-lactams were synthesized in two steps from commercially available 3-bromo-2-(bromomethyl)propionic acid and reacted with arene diazonium salts in a Heck-type arylation in the presence of catalytic amounts of Pd(OAc)2 under ligand-free conditions. The products, arylidene-ß-lactams, were obtained in high yields as single isomers. The ß-hydride elimination step of the Pd-catalyzed coupling reaction proceeds with high exo-regioselectivity and E-stereoselectivity. With aryl iodides, triflates, or bromides, the coupling products were isolated only in low yields, due to extensive decomposition of the starting material at elevated temperatures. This underlines that arene diazonium salts can be superior arylating reagents in Heck-type reactions and yield coupling products in synthetically useful yields and selectivities when conventional conditions fail.

Age-related changes in time perception: The impact of naturalistic environments and retrospective judgements on timing performance.

Reduced timing abilities have been reported in older adults and are associated with pathological cognitive decline. However, time perception experiments often lack ecological validity. Especially the reduced complexity of experimental stimuli and the participants' awareness of the time-related nature of the task can influence lab-assessed timing performance and thereby conceal age-related differences. An approximation of more naturalistic paradigms can provide important information about age-related changes in timing abilities. To determine the impact of higher ecological validity on timing experiments, we implemented a paradigm that allowed us to test (1) the effect of embedding the to-be-timed stimuli within a naturalistic visual scene and (2) the effect of retrospective time judgements, which are more common in real life than prospective judgements. The results show that compared with out-of-context stimuli, younger adults benefit from a naturalistic embedding of stimuli (reflected in higher precision and less errors), whereas the performance of older adults is reduced when confronted with naturalistic stimuli. Differences between retrospective and prospective time judgements were not modulated by age. We conclude that, potentially driven by difficulties in suppressing temporally irrelevant environmental information, the contextual embedding of naturalistic stimuli can affect the degree to which age influences the performance in time perception tasks.

Spatial Updating Depends on Gravity.

As we move through an environment the positions of surrounding objects relative to our body constantly change. Maintaining orientation requires spatial updating, the continuous monitoring of self-motion cues to update external locations. This ability critically depends on the integration of visual, proprioceptive, kinesthetic, and vestibular information. During weightlessness gravity no longer acts as an essential reference, creating a discrepancy between vestibular, visual and sensorimotor signals. Here, we explore the effects of repeated bouts of microgravity and hypergravity on spatial updating performance during parabolic flight. Ten healthy participants (four women, six men) took part in a parabolic flight campaign that comprised a total of 31 parabolas. Each parabola created about 20-25 s of 0 g, preceded and followed by about 20 s of hypergravity (1.8 g). Participants performed a visual-spatial updating task in seated position during 15 parabolas. The task included two updating conditions simulating virtual forward movements of different lengths (short and long), and a static condition with no movement that served as a control condition. Two trials were performed during each phase of the parabola, i.e., at 1 g before the start of the parabola, at 1.8 g during the acceleration phase of the parabola, and during 0 g. Our data demonstrate that 0 g and 1.8 g impaired pointing performance for long updating trials as indicated by increased variability of pointing errors compared to 1 g. In contrast, we found no support for any changes for short updating and static conditions, suggesting that a certain degree of task complexity is required to affect pointing errors. These findings are important for operational requirements during spaceflight because spatial updating is pivotal for navigation when vision is poor or unreliable and objects go out of sight, for example during extravehicular activities in space or the exploration of unfamiliar environments. Future studies should compare the effects on spatial updating during seated and free-floating conditions, and determine at which g-threshold decrements in spatial updating performance emerge.

Sources of path integration error in young and aging humans.

Path integration plays a vital role in navigation: it enables the continuous tracking of one's position in space by integrating self-motion cues. Path integration abilities vary widely across individuals, and tend to deteriorate in old age. The specific causes of path integration errors, however, remain poorly characterized. Here, we combine tests of path integration performance in participants of different ages with an analysis based on the Langevin equation for diffusive dynamics, which allows us to decompose errors into distinct causes that can corrupt path integration computations. We show that, across age groups, the dominant error source is unbiased noise that accumulates with travel distance not elapsed time, suggesting that the noise originates in the velocity input rather than within the integrator. Age-related declines are primarily traced to a growth in this noise. These findings shed light on the contributors to path integration error and the mechanisms underlying age-related navigational deficits.

Which Effects on Neuroanatomy and Path-Integration Survive? Results of a Randomized Controlled Study on Intensive Balance Training.

Balancing is a complex task requiring the integration of visual, somatosensory and vestibular inputs. The vestibular system is linked to the hippocampus, a brain structure crucial for spatial orientation. Here we tested the immediate and sustained effects of a one-month-long slackline training program on balancing and orientation abilities as well as on brain volumes in young adults without any prior experience in that skill. On the corrected level, we could not find any interaction effects for brain volumes, but the effect sizes were small to medium. A subsequent within-training-group analysis revealed volumetric increments within the somatosensory cortex and decrements within posterior insula, cerebellum and putamen remained stable over time. No significant interaction effects were observed on the clinical balance and the spatial orientation task two months after the training period (follow-up). We interpret these findings as a shift away from processes crucial for automatized motor output towards processes related to voluntarily controlled movements. The decrease in insular volume in the training group we propose to result from multisensory interaction of the vestibular with the visual and somatosensory systems. The discrepancy between sustained effects in the brain of the training group on the one hand and transient benefits in function on the other may indicate that for the latter to be retained a longer-term practice is required.

Negative errors in time reproduction tasks.

In time reproduction tasks, the reaction time of motor responses is intrinsically linked to the measure of perceptual timing. Decisions are based on a continuous comparison between elapsed time and a memory trace of the to-be-reproduced interval. Here, we investigate the possibility that negative reproduction errors can be explained by the tendency to prefer earlier over later response times, or whether the whole range of possible response times is shifted. In experiment 1, we directly compared point reproduction (participants indicate the exact time point of equality) and range reproduction (participants bracket an interval containing this time point). In experiment 2, participants indicated, in three separate tasks, the exact time point at which the reproduction phase was equal to the standard duration (point reproduction), the earliest (start reproduction), or the latest moment (stop reproduction) at which the exact time point of equality might have been reached. The results demonstrate that the bias towards earlier responses not only affects reproduction of the exact time point of equality. Rather, even if the decision threshold is changed in favor of late responses, they exhibit a continuous shift towards negative errors that increases with the length of the standard duration. The findings are discussed in the context of the hypothesis that systematic errors in time reproduction tasks reflect a dimension-unspecific tendency towards earlier responses caused by the psychophysical method rather than by a time-specific perceptual distortion.

Reversing Threat to Safety: Incongruence of Facial Emotions and Instructed Threat Modulates Conscious Perception but Not Physiological Responding.

Facial expressions inform about other peoples' emotion and motivation and thus are central for social communication. However, the meaning of facial expressions may change depending on what we have learned about the related consequences. For instance, a smile might easily become threatening when displayed by a person who is known to be dangerous. The present study examined the malleability of emotional facial valence by means of social learning. To this end, facial expressions served as cues for verbally instructed threat-of-shock or safety (e.g., "happy faces cue shocks"). Moreover, reversal instructions tested the flexibility of threat/safety associations (e.g., "now happy faces cue safety"). Throughout the experiment, happy, neutral, and angry facial expressions were presented and auditory startle probes elicited defensive reflex activity. Results show that self-reported ratings and physiological reactions to threat/safety cues dissociate. Regarding threat and valence ratings, happy facial expressions tended to be more resistant becoming a threat cue, and angry faces remain threatening even when instructed as safety cue. For physiological response systems, however, we observed threat-potentiated startle reflex and enhanced skin conductance responses for threat compared to safety cues regardless of whether threat was cued by happy or angry faces. Thus, the incongruity of visual and verbal threat/safety information modulates conscious perception, but not the activation of physiological response systems. These results show that verbal instructions can readily overwrite the intrinsic meaning of facial emotions, with clear benefits for social communication as learning and anticipation of threat and safety readjusted to accurately track environmental changes.

Timing deficiencies in amnestic Mild Cognitive Impairment: Disentangling clock and memory processes.

Interval timing performance in cognitive decline is typically characterized by decreased accuracy, precision, or both. One explanation for this decreased performance is a larger clock time variability. However, memory deficiencies associated with cognitive decline might also affect temporal performance in two alternative ways: First, memory deficiencies could lead to reduced encoding of just perceived durations, and thus a stronger reliance on the memory traces of previous experiences (the "prior"), yielding less precise reproductions of the most current experiences. Second, memory deficiencies could hamper the storage of perceived intervals, thus resulting in less influence of the prior. Here, we present data of 15 patients with amnestic Mild Cognitive Impairment (aMCI) and 44 healthy, aged controls, the latter split in two groups based on memory performance. All participants were tested on a temporal production task to assess clock time variability and a multi-duration reproduction task to assess the influence of memory traces reflecting current and previous experiences. Patients with aMCI showed the strongest regression towards the mean in a multi-duration reproduction task, followed by low-performing healthy controls and high-performing healthy controls, respectively. As no difference was observed between the groups in terms of clock time variability, and clock variability did not statistically contribute to the observed regression, this increased central tendency effect was not attributable to clock noise. We therefore, in line with the first explanation, conclude that memory deficiencies result in a stronger (relative) reliance on the prior.

The rubber hand universe: On the impact of methodological differences in the rubber hand illusion.

The rubber hand illusion (RHI) is a widely applied paradigm to investigate changes in body representations. Extensive scientific interest has produced a great variability in the observed results and many contradictory findings have been reported. Taking into account the numerous variations in the experimental implementation of the RHI, many of these contradictive findings can be reconciled, but to date a thorough analysis of the methodological differences between RHI studies is lacking. Here we summarize and analyse methodological differences between RHI studies. In distinction from other reviews focusing on the integration of findings from various studies, the present paper is devoted to the differences in (i) the experimental setup, (ii) the method used to induce the RHI, (iii) the quantification of its effects, and (iv) aspects of the experimental design and data analysis. This approach will provide a reference frame for the interpretation of previous studies as well as for the design of future studies.

The effect of feedback on temporal error monitoring and timing behavior.

Metacognitive processes in human timing behavior are rarely investigated, which stands in sharp contrast to the wide research field of metacognition itself. To date, little is known about the sources and the reliability of information that humans possess to judge their own timing abilities and to monitor errors in time-keeping. Here, we intended to fill this gap by determining the degree to which humans depend on external feedback to adjust their timing behavior and make metacognitive accuracy judgments. Two groups of participants performed a time reproduction task under different feedback conditions. After each trial, participants were informed either about the magnitude and the direction of their timing error (signed feedback group) or about its magnitude alone (absolute feedback group). Reproduction errors were related to retrospective, metacognitive judgments on the overall timing performance. The results indicate that the under reproduction effect occurred, rather independently of the type of feedback; however, signed feedback was essential to reduce the bias in metacognitive judgments on timing accuracy. Without being explicitly informed about the direction of timing errors (whether the reproduction interval was stopped too early or too late), participants significantly overestimated their reproduced durations. These results extend previous reports of metacognitive processes in timing behavior measured on a single-trial basis, and provide new insights into the ability of temporal error monitoring in humans.

Preserved multisensory body representations in advanced age.

The internal representation of the body emerges via the integration of multisensory body cues. Sensory signal transfer and the ability to integrate multisensory information deteriorate significantly with increasing age. However, there is little empirical evidence on age-related changes in body representations based on multisensory integration. Here, we used a standard paradigm for evaluating body representations based on multisensory integration, the rubber hand illusion, and compared the amount of proprioceptive drift and changes in perceived body ownership triggered by the integration of visual, tactile, and proprioceptive cues between younger and older adults. To account for potential age-related differences in the temporal stability of the illusion, proprioceptive drift was measured at five different time points. Our results show that older adults used synchronous visuo-tactile cues similarly to younger adults to update both the position of their own hand, and their feeling of ownership over the artificial hand. Independent of visuo-tactile synchrony, older adults perceived their hand as closer to their body than younger adults did, and showed a less stable representation of this in-depth hand position. This proprioceptive bias towards the body did not correlate with the strength of the illusion. Our results indicate that the integration of visual and tactile cues is largely preserved in advanced age when used to update limb position, whereas proprioception worsens with age. This may be linked to two different pathways that underlie changes in body representations over the life span.