Age effects on predictive eye movements for action.

When interacting with the environment, humans typically shift their gaze to where information is to be found that is useful for the upcoming action. With increasing age, people become slower both in processing sensory information and in performing their movements. One way to compensate for this slowing down could be to rely more on predictive strategies. To examine whether we could find evidence for this, we asked younger (19-29 years) and older (55-72 years) healthy adults to perform a reaching task wherein they hit a visual target that appeared at one of two possible locations. In separate blocks of trials, the target could appear always at the same location (predictable), mainly at one of the locations (biased), or at either location randomly (unpredictable). As one might expect, saccades toward predictable targets had shorter latencies than those toward less predictable targets, irrespective of age. Older adults took longer to initiate saccades toward the target location than younger adults, even when the likely target location could be deduced. Thus we found no evidence of them relying more on predictive gaze. Moreover, both younger and older participants performed more saccades when the target location was less predictable, but again no age-related differences were found. Thus we found no tendency for older adults to rely more on prediction.

Aging attenuates the memory advantage for unexpected objects in real-world scenes.

Across the adult lifespan memory processes are subject to pronounced changes. Prior knowledge and expectations might critically shape functional differences; however, corresponding findings have remained ambiguous so far. Here, we chose a tailored approach to scrutinize how schema (in-)congruencies affect older and younger adults' memory for objects embedded in real-world scenes, a scenario close to everyday life memory demands. A sample of 23 older (52-81 years) and 23 younger adults (18-38 years) freely viewed 60 photographs of scenes in which target objects were included that were either congruent or incongruent with the given context information. After a delay, recognition performance for those objects was determined. In addition, recognized objects had to be matched to the scene context in which they were previously presented. While we found schema violations beneficial for object recognition across age groups, the advantage was significantly less pronounced in older adults. We moreover observed an age-related congruency bias for matching objects to their original scene context. Our findings support a critical role of predictive processes for age-related memory differences and indicate enhanced weighting of predictions with age. We suggest that recent predictive processing theories provide a particularly useful framework to elaborate on age-related functional vulnerabilities as well as stability.

Cross-modal metacognition: Visual and tactile confidence share a common scale.

Humans can judge the quality of their perceptual decisions-an ability known as perceptual confidence. Previous work suggested that confidence can be evaluated on an abstract scale that can be sensory modality-independent or even domain-general. However, evidence is still scarce on whether confidence judgments can be directly made across visual and tactile decisions. Here, we investigated in a sample of 56 adults whether visual and tactile confidence share a common scale by measuring visual contrast and vibrotactile discrimination thresholds in a confidence-forced choice paradigm. Confidence judgments were made about the correctness of the perceptual decision between two trials involving either the same or different modalities. To estimate confidence efficiency, we compared discrimination thresholds obtained from all trials to those from trials judged to be relatively more confident. We found evidence for metaperception because higher confidence was associated with better perceptual performance in both modalities. Importantly, participants were able to judge their confidence across modalities without any costs in metaperceptual sensitivity and only minor changes in response times compared to unimodal confidence judgments. In addition, we were able to predict cross-modal confidence well from unimodal judgments. In conclusion, our findings show that perceptual confidence is computed on an abstract scale and that it can assess the quality of our decisions across sensory modalities.

Age-related differences in visual confidence are driven by individual differences in cognitive control capacities.

Visual perception is not only shaped by sensitivity but also by confidence, i.e., the ability to estimate the accuracy of a visual decision. Younger observers have been reported to have access to a reliable measure of their own uncertainty when making visual decisions. This metacognitive ability might be challenged during ageing due to increasing sensory noise and decreasing cognitive control resources. We investigated age effects on visual confidence using a visual contrast discrimination task and a confidence forced-choice paradigm. Younger adults (19-38 years) showed significantly lower discrimination thresholds than older adults (60-78 years). To focus on confidence sensitivity above and beyond differences in discrimination performance, we estimated confidence efficiency that reflects the ability to distinguish good from bad perceptual decisions. Confidence efficiency was estimated by comparing thresholds obtained from all trials and trials that were judged with relatively higher confidence, respectively. In both age groups, high confidence judgments were associated with better visual performance, but confidence efficiency was reduced in older adults. However, we observed substantial variability across all participants. Controlling for age group, confidence effciency was closely linked to individual differences in cognitive control capacities. Our findings provide evidence for age-related differences in confidence efficiency that present a specific challenge to perceptual performance in old age. We propose that these differences are driven by cognitive control capacities, supporting their crucial role for metacognitive efficiency.

Age-related effects on the neural processing of semantic complexity in a continuous narrative: Modulation by gestures already present in young to middle-aged adults.

The processing of semantically complex speech is a demanding task which can be facilitated by speech-associated arm and hand gestures. However, the role of age concerning the perception of semantic complexity and the influence of gestures in this context remains unclear. The goal of this study was to investigate if age-related differences are already present in early adulthood during the processing of semantic complexity and gestures. To this end, we analyzed fMRI images of a sample of 38 young and middle-aged participants (age-range: 19-55). They had the task to listen and to watch a narrative. The narrative contained segments varying in the degree of semantic complexity, and they were spontaneously accompanied by gestures. The semantic complexity of the story was measured by the idea density. Consistent with previous findings in young adults, we observed increased activation for passages with lower compared to higher complexity in bilateral temporal areas and the precuneus. BOLD signal in the left frontal and left parietal regions correlated during the perception of complex passages with increasing age. This correlation was reduced for passages presented with gestures. Median-split based post-hoc comparisons confirmed that group differences between younger (19-23 years) and older adults within the early adult lifespan (24-55 years) were significantly reduced in passages with gestures. Our results suggest that older adults within early adulthood adapt to the requirements of highly complex passages activating additional regions when no gesture information is available. Gestures might play a facilitative role with increasing age, especially when speech is complex.

Individual differences in processing resources modulate bimanual interference in pointing.

Coordinating both hands during bimanual reaching is a complex task that can generate interference during action preparation as often indicated by prolonged reaction times for movements that require moving the two hands at different amplitudes. Individual processing constraints are thought to contribute to this interference effect. Most importantly, however, the amount of interference seems to depend considerably on overall task demands suggesting that interference increases as the available processing resources decrease. Here, we further investigated this idea by comparing performance in a simple direct cueing and a more difficult symbolic cueing task between three groups of participants that supposedly vary in their processing resources, i.e., musicians, young adults and older adults. We found that the size of interference effects during symbolic cueing varied in the tested groups: musicians showed the smallest and older adults the largest interference effects. More importantly, a regression model, using processing speed and processing capacity as predictor variables, revealed a clear link between the available processing resources and the size of the interference effect during symbolic cueing. In the easier direct cueing task, no reliable interference was observed on a group level. We propose that the susceptibility to bimanual interference is modulated by the task-specific processing requirements in relation with the available processing resources of an individual.

Age effects on sensorimotor predictions: What drives increased tactile suppression during reaching?

Tactile suppression refers to the phenomenon that tactile signals are attenuated during movement planning and execution when presented on a moving limb compared to rest. It is usually explained in the context of the forward model of movement control that predicts the sensory consequences of an action. Recent research suggests that aging increases reliance on sensorimotor predictions resulting in stronger somatosensory suppression. However, the mechanisms contributing to this age effect remain to be clarified. We measured age-related differences in tactile suppression during reaching and investigated the modulation by cognitive processes. A total of 23 younger (18-27 years) and 26 older (59-78 years) adults participated in our study. We found robust suppression of tactile signals when executing reaching movements. Age group differences corroborated stronger suppression in old age. Cognitive task demands during reaching, although overall boosting suppression effects, did not modulate the age effect. Across age groups, stronger suppression was associated with lower individual executive capacities. There was no evidence that baseline sensitivity had a prominent impact on the magnitude of suppression. We conclude that aging alters the weighting of sensory signals and sensorimotor predictions during movement control. Our findings suggest that individual differences in tactile suppression are critically driven by executive functions.

Motion perception as a model for perceptual aging.

Research on functional changes across the adult lifespan has been dominated by studies related to cognitive processes. However, it has become evident that a more comprehensive approach to behavioral aging is needed. In particular, our understanding of age-related perceptual changes is limited. Visual motion perception is one of the most studied areas in perceptual aging and therefore, provides an excellent domain on the basis of which we can investigate the complexity of the aging process. We review the existing literature on how aging affects motion perception, including different processing stages, and consider links to cognitive and motor changes. We address the heterogeneity of results and emphasize the role of individual differences. Findings on age-related changes in motion perception ultimately illustrate the complexity of functional dynamics that can contribute to decline as well as stability during healthy aging. We thus propose that motion perception offers a conceptual framework for perceptual aging, encouraging a deliberate consideration of functional limits and resources emerging across the lifespan.

Motivational Modulation of Age-Related Effects on Reaching Adaptation.

Previous studies have provided consistent evidence that adaptation to visuomotor rotations during reaching declines with age. Since it has been recently shown that learning and retention components of motor adaptation are modulated by reward and punishment, we were interested in how motivational feedback affects age-related decline in reaching adaptation. We studied 35 young and 32 older adults in a reaching task which required fast shooting movements toward visual targets with their right hand. A robotic manipulandum (vBOT system) allowed measuring reaching trajectories. Targets and visual feedback on hand position were presented using a setup that prevented direct vision of the hand and projected a virtual image by a semi-silvered mirror. After a baseline block with veridical visual feedback we introduced a 30° counterclockwise visuomotor rotation. After this adaptation block we also measured retention of adaptation without visual feedback and finally readaptation for the previously experienced rotation. In the adaptation block participants were assigned to one of three motivational feedback conditions, i.e., neutral, reward, or punishment. Reward and punishment feedback was based on reaching endpoint error. Our results consistently corroborated reduced motor learning capacities in older adults (p < 0.001, η2 = 0.56). However, motivational feedback modulated learning rates equivalently in both age groups (p = 0.028, η2 = 0.14). Rewarding feedback induced faster learning, though punishing feedback had no effect. For retention we determined a significant interaction effect between motivational feedback and age group (p = 0.032, η2 = 0.13). Previously provided motivational feedback was detrimental for young adults, but not for older adults. We did not observe robust effects of motivational feedback on readaptation (p = 0.167, η2 = 0.07). Our findings support that motor learning is subject to modulation by motivational feedback. Whereas learning is boosted across both age groups, retention is vulnerable to previously experienced motivational incentives in young adults. In summary, in particular older adults benefit from motivational feedback during reaching adaptation so that age-related differences in visuomotor plasticity, though persisting, can be attenuated. We suggest that the use of motivational information provides a potentially compensatory mechanism during functional aging.

Avoiding unseen obstacles: Subcortical vision is not sufficient to maintain normal obstacle avoidance behaviour during reaching.

Previous research found that a patient with cortical blindness (homonymous hemianopia) was able to successfully avoid an obstacle placed in his blind field, despite reporting no conscious awareness of it [Striemer, C. L., Chapman, C. S., & Goodale, M. A., 2009, PNAS, 106(37), 15996-16001]. This finding led to the suggestion that dorsal stream areas, that are assumed to mediate obstacle avoidance behaviour, may obtain their visual input primarily from subcortical pathways. Hence, it was suggested that normal obstacle avoidance behaviour can proceed without input from the primary visual cortex. Here we tried to replicate this finding in a group of patients (N = 6) that suffered from highly circumscribed lesions in the occipital lobe (including V1) that spared the subcortical structures that have been associated with action-blindsight. We also tested if obstacle avoidance behaviour differs depending on whether obstacles are placed only in the blind field or in both the blind and intact visual field of the patients simultaneously. As expected, all patients successfully avoided obstacles placed in their intact visual field. However, none of them showed reliable avoidance behaviour - as indicated by adjustments in the hand trajectory in response to obstacle position - for obstacles placed in their blind visual field. The effects were not dependent on whether one or two obstacles were present. These findings suggest that behaviour in complex visuomotor tasks relies on visual input from occipital areas.

Healthy aging is associated with decreased risk-taking in motor decision-making.

Healthy aging is associated with changes in both cognitive abilities, including decision-making, and motor control. Previous research has shown that young healthy observers are close to optimal when they perform a motor equivalent of economic decision-making tasks that are known to produce suboptimal decision patterns. We tested both younger (age 20-29) and older (age 60-79) adults in such a task, which involved rapid manual aiming and monetary rewards and punishments contingent on hitting different areas on a touch screen. Older adults were as close to optimal as younger adults at the task, but differed from the younger adults in their strategy. Older adults appeared to be relatively less risk-seeking, as evidenced by the fact that they adjusted their aiming strategy to a larger extent to avoid the penalty area. A model-based interpretation of the results suggested that the change in aiming strategy between younger and older adults was mainly driven by the fact that the first weighted monetary gains more than losses, rather than by a mis-estimation of one's motor variability. The results parallel the general finding that older adults tend to be less risk-seeking than younger adults in economic decision-making and complement the observation that children are even more risk-seeking than younger adults in a similar motor decision-making paradigm. (PsycINFO Database Record

Age Effects on Visuo-Haptic Length Discrimination: Evidence for Optimal Integration of Senses in Senior Adults.

Demographic changes in most developed societies have fostered research on functional aging. While cognitive changes have been characterized elaborately, understanding of perceptual aging lacks behind. We investigated age effects on the mechanisms of how multiple sources of sensory information are merged into a common percept. We studied visuo-haptic integration in a length discrimination task. A total of 24 young (20-25 years) and 27 senior (69-77 years) adults compared standard stimuli to appropriate sets of comparison stimuli. Standard stimuli were explored under visual, haptic, or visuo-haptic conditions. The task procedure allowed introducing an intersensory conflict by anamorphic lenses. Comparison stimuli were exclusively explored haptically. We derived psychometric functions for each condition, determining points of subjective equality and discrimination thresholds. We notably evaluated visuo-haptic perception by different models of multisensory processing, i.e., the Maximum-Likelihood-Estimate model of optimal cue integration, a suboptimal integration model, and a cue switching model. Our results support robust visuo-haptic integration across the adult lifespan. We found suboptimal weighted averaging of sensory sources in young adults, however, senior adults exploited differential sensory reliabilities more efficiently to optimize thresholds. Indeed, evaluation of the MLE model indicates that young adults underweighted visual cues by more than 30%; in contrast, visual weights of senior adults deviated only by about 3% from predictions. We suggest that close to optimal multisensory integration might contribute to successful compensation for age-related sensory losses and provides a critical resource. Differentiation between multisensory integration during healthy aging and age-related pathological challenges on the sensory systems awaits further exploration.

Links between Gestures and Multisensory Processing: Individual Differences Suggest a Compensation Mechanism.

Speech-associated gestures represent an important communication modality. However, individual differences in the production and perception of gestures are not well understood so far. We hypothesized that the perception of multisensory action consequences might play a crucial role. Verbal communication involves continuous calibration of audio-visual information produced by the speakers. The effective production and perception of gestures supporting this process could depend on the given capacities to perceive multisensory information accurately. We explored the association between the production and perception of gestures and the monitoring of multisensory action consequences in a sample of 31 participants. We applied a recently introduced gesture scale to assess self-reported gesture production and perception in everyday life situations. In the perceptual experiment, we presented unimodal (visual) and bimodal (visual and auditory) sensory outcomes with various delays after a self-initiated (active) or externally generated (passive) button press. Participants had to report whether they detected a delay between the button press and the visual stimulus. We derived psychometric functions for each condition and determined points of subjective equality, reflecting detection thresholds for delays. Results support a robust link between gesture scores and detection thresholds. Individuals with higher detection thresholds (lower performance) reported more frequent gesture production and perception and furthermore profited more from multisensory information in the experimental task. We propose that our findings indicate a compensational function of multisensory processing as a basis for individual differences in both action outcome monitoring and gesture production and perception in everyday life situations.

Age effects on saccadic adaptation: Evidence from different paradigms reveals specific vulnerabilities.

Saccadic eye movements provide an opportunity to study closely interwoven perceptual, motor, and cognitive changes during aging. Here, we investigated age effects on different mechanisms of saccadic plasticity. We compared age effects in two different adaptation paradigms that tap into low- and high-level adaptation processes. A total of 27 senior adults and 25 young adults participated in our experiments. In our first experiment, we elicited adaptation by a double-step paradigm, which is designed to trigger primarily low-level, gradual motor adaptation. Age groups showed equivalent adaptation of saccadic gain. In our second experiment, adaptation was induced by a perceptual task that emphasizes high-level, fast processes. We consistently found no evidence for age-related differences in low-level adaptation; however, the fast adaptation response was significantly more pronounced in the young adult group. We conclude that low-level motor adaptation is robust during healthy aging but that high-level contributions, presumably involving executive strategies, are subject to age-related decline. Our findings emphasize the need to differentiate between specific aging processes in order to understand functional decline and stability across the adult life span.

Keeping Safe: Intra-individual Consistency in Obstacle Avoidance Behaviour Across Grasping and Locomotion Tasks.

Successful obstacle avoidance requires a close coordination of the visual and the motor systems. Visual information is essential for adjusting movements in order to avoid unwanted collisions. Yet, established obstacle avoidance paradigms have typically either focused on gaze strategies or on motor adjustments. Here we were interested in whether humans show similar visuomotor sensitivity to obstacles when gaze and motor behaviour are measured across different obstacle avoidance tasks. To this end, we measured participants' hand movement paths when grasping targets in the presence of obstacles as well as their gaze behaviour when walking through a cluttered hallway. We found that participants who showed more pronounced motor adjustments during grasping also spent more time looking at obstacles during locomotion. Furthermore, movement durations correlated positively in both tasks. Results suggest considerable intra-individual consistency in the strength of the avoidance response across different visuomotor measures potentially indicating an individual's tendency to perform safe actions.

Association between COMT genotype and the control of memory guided saccades: Individual differences in healthy adults reveal a detrimental role of dopamine.

The neural circuits involved in oculomotor control are well described; however, neuromodulation of eye movements is still hardly understood. Memory guided saccades have been extensively studied and in particular neurophysiological evidence from monkey studies points to a crucial functional role of prefrontal dopamine activity. We exploited individual differences in dopamine regulation due to the well established COMT (catechol-O-methyltransferase) Val158Met polymorphism to explore the link between prefrontal dopamine activity and memory guided saccades in healthy subjects. The COMT genotype is thought to modulate dopamine metabolism in prefrontal cortex producing differences in dopamine availability. We investigated memory guided saccades in 111 healthy subjects and determined individual genotypes. Accuracy and precision were reduced in subjects with putatively higher prefrontal dopamine levels. In contrast, we found no modulation of saccade parameters by genotype in a visually guided control task. Our results suggest that increased dopamine activity can have a detrimental effect on saccades that rely on spatial memory representations. Although these findings await replication in larger and more diverse sample sizes, they provide persuasive support that specific oculomotor parameters are sensitive to dopaminergic variation in healthy subjects and add to a better understanding of how dopamine modulates saccadic control.

The Role of Dopamine in Anticipatory Pursuit Eye Movements: Insights from Genetic Polymorphisms in Healthy Adults.

There is a long history of eye movement research in patients with psychiatric diseases for which dysfunctions of neurotransmission are considered to be the major pathologic mechanism. However, neuromodulation of oculomotor control is still hardly understood. We aimed to investigate in particular the impact of dopamine on smooth pursuit eye movements. Systematic variability in dopaminergic transmission due to genetic polymorphisms in healthy subjects offers a noninvasive opportunity to determine functional associations. We measured smooth pursuit in 110 healthy subjects genotyped for two well-documented polymorphisms, the COMT Val158Met polymorphism and the SLC6A3 3'-UTR-VNTR polymorphism. Pursuit paradigms were chosen to particularly assess the ability of the pursuit system to initiate tracking when target motion onset is blanked, reflecting the impact of extraretinal signals. In contrast, when following a fully visible target sensory, retinal signals are available. Our results highlight the crucial functional role of dopamine for anticipatory, but not for sensory-driven, pursuit processes. We found the COMT Val158Met polymorphism specifically associated with anticipatory pursuit parameters, emphasizing the dominant impact of prefrontal dopamine activity on complex oculomotor control. In contrast, modulation of striatal dopamine activity by the SLC6A3 3'-UTR-VNTR polymorphism had no significant functional effect. Though often neglected so far, individual differences in healthy subjects provide a promising approach to uncovering functional mechanisms and can be used as a bridge to understanding deficits in patients.

Robust Underestimation of Speed During Driving: A Field Study.

Traffic reports consistently identify speeding as a substantial source of accidents. Adequate driving speeds require reliable speed estimation; however, there is still a lack of understanding how speed perception is biased during driving. Therefore, we ran three experiments measuring speed estimation under controlled driving and lighting conditions. In the first experiment, participants had to produce target speeds as drivers or had to judge driven speed as passengers. Measurements were performed at daylight and at night. In the second experiment, participants were required to produce target speeds at dusk, under rapidly changing lighting conditions. In the third experiment, we let two cars approach and pass each other. Drivers were instructed to produce target speeds as well as to judge the speed of the oncoming vehicle. Here measurements were performed at daylight and at night, with full or dipped headlights. We found that passengers underestimated driven speed by about 20% and drivers went over the instructed speed by roughly the same amount. Interestingly, the underestimation of speed extended to oncoming cars. All of these effects were independent of lighting conditions. The consistent underestimation of speed could lead to potentially fatal situations where drivers go faster than intended and judge oncoming traffic to approach slower than it actually is.

Challenges to normal neural functioning provide insights into separability of motion processing mechanisms.

There is a long history of attempts to disentangle different visual processing mechanisms for physically different motion cues. However, underlying neural correlates and separability of networks are still under debate. We aimed to refine the current understanding by studying differential vulnerabilities when normal neural functioning is challenged. We investigated effects of ageing and extrastriate brain lesions on detection thresholds for motion defined by either luminance- or contrast modulations, known as first- and second-order motion. Both approaches focus on extrastriate processing changes and combine distributed as well as more focal constraints. Our ageing sample comprised 102 subjects covering an age range from 20 to 82 years. Threshold signal-to-noise ratios for detection approximately doubled across the age range for both motion types. Results suggest that ageing affects perception of both motion types to an equivalent degree and thus support overlapping processing resources. Underlying neural substrates were further qualified by testing perceptual performance of 18 patients with focal cortical brain lesions. We determined selective first-order motion deficits in three patients, selective second-order motion deficits in only one patient, and deficits for both motion types in three patients. Lesion analysis yielded support for common functional substrates in higher cortical regions. Functionally specific substrates remained ambiguous, but tended to cover earlier visual areas. We conclude that observed vulnerabilities of first- and second-order motion perception provide limited evidence for functional specialization at early extrastriate stages, but emphasize shared processing pathways at higher cortical levels.