Multiplicative joint coding in preparatory activity for reaching sequence in macaque motor cortex.

Although the motor cortex has been found to be modulated by sensory or cognitive sequences, the linkage between multiple movement elements and sequence-related responses is not yet understood. Here, we recorded neuronal activity from the motor cortex with implanted micro-electrode arrays and single electrodes while monkeys performed a double-reach task that was instructed by simultaneously presented memorized cues. We found that there existed a substantial multiplicative component jointly tuned to impending and subsequent reaches during preparation, then the coding mechanism transferred to an additive manner during execution. This multiplicative joint coding, which also spontaneously emerged in recurrent neural networks trained for double reach, enriches neural patterns for sequential movement, and might explain the linear readout of elemental movements.

A bonus task boosts people's willingness to offload cognition to an algorithm.

With the increased sophistication of technology, humans have the possibility to offload a variety of tasks to algorithms. Here, we investigated whether the extent to which people are willing to offload an attentionally demanding task to an algorithm is modulated by the availability of a bonus task and by the knowledge about the algorithm's capacity. Participants performed a multiple object tracking (MOT) task which required them to visually track targets on a screen. Participants could offload an unlimited number of targets to a "computer partner". If participants decided to offload the entire task to the computer, they could instead perform a bonus task which resulted in additional financial gain-however, this gain was conditional on a high performance accuracy in the MOT task. Thus, participants should only offload the entire task if they trusted the computer to perform accurately. We found that participants were significantly more willing to completely offload the task if they were informed beforehand that the computer's accuracy was flawless (Experiment 1 vs. 2). Participants' offloading behavior was not significantly affected by whether the bonus task was incentivized or not (Experiment 2 vs. 3). These results combined with those from our previous study (Wahn et al. in PLoS ONE 18:e0286102, 2023), which did not include a bonus task but was identical otherwise, show that the human willingness to offload an attentionally demanding task to an algorithm is considerably boosted by the availability of a bonus task-even if not incentivized-and by the knowledge about the algorithm's capacity.

Position- and scale-invariant object-centered spatial localization in monkey frontoparietal cortex dynamically adapts to cognitive demand.

Egocentric encoding is a well-known property of brain areas along the dorsal pathway. Different to previous experiments, which typically only demanded egocentric spatial processing during movement preparation, we designed a task where two male rhesus monkeys memorized an on-the-object target position and then planned a reach to this position after the object re-occurred at variable location with potentially different size. We found allocentric (in addition to egocentric) encoding in the dorsal stream reach planning areas, parietal reach region and dorsal premotor cortex, which is invariant with respect to the position, and, remarkably, also the size of the object. The dynamic adjustment from predominantly allocentric encoding during visual memory to predominantly egocentric during reach planning in the same brain areas and often the same neurons, suggests that the prevailing frame of reference is less a question of brain area or processing stream, but more of the cognitive demands.

Perturbation Variability Does Not Influence Implicit Sensorimotor Adaptation.

Implicit adaptation has been regarded as a rigid process that automatically operates in response to movement errors to keep the sensorimotor system precisely calibrated. This hypothesis has been challenged by recent evidence suggesting flexibility in this learning process. One compelling line of evidence comes from work suggesting that this form of learning is context-dependent, with the rate of learning modulated by error history. Specifically, learning was attenuated in the presence of perturbations exhibiting high variance compared to when the perturbation is fixed. However, these findings are confounded by the fact that the adaptation system corrects for errors of different magnitudes in a non-linear manner, with the adaptive response increasing in a proportional manner to small errors and saturating to large errors. Through simulations, we show that this non-linear motor correction function is sufficient to explain the effect of perturbation variance without referring to an experience-dependent change in error sensitivity. Moreover, by controlling the distribution of errors experienced during training, we provide empirical evidence showing that there is no measurable effect of perturbation variance on implicit adaptation. As such, we argue that the evidence to date remains consistent with the rigidity assumption.

Evaluation of power wheelchair driving performance in simulator compared to driving in real-life situations: the SIMADAPT (simulator ADAPT) project-a pilot study.

OBJECTIVE: The objective of this study was to evaluate users' driving performances with a Power Wheelchair (PWC) driving simulator in comparison to the same driving task in real conditions with a standard power wheelchair. METHODS: Three driving circuits of progressive difficulty levels (C1, C2, C3) that were elaborated to assess the driving performances with PWC in indoor situations, were used in this study. These circuits have been modeled in a 3D Virtual Environment to replicate the three driving task scenarios in Virtual Reality (VR). Users were asked to complete the three circuits with respect to two testing conditions during three successive sessions, i.e. in VR and on a real circuit (R). During each session, users completed the two conditions. Driving performances were evaluated using the number of collisions and time to complete the circuit. In addition, driving ability by Wheelchair Skill Test (WST) and mental load were assessed in both conditions. Cybersickness, user satisfaction and sense of presence were measured in VR. The conditions R and VR were randomized. RESULTS: Thirty-one participants with neurological disorders and expert wheelchair drivers were included in the study. The driving performances between VR and R conditions were statistically different for the C3 circuit but were not statistically different for the two easiest circuits C1 and C2. The results of the WST was not statistically different in C1, C2 and C3. The mental load was higher in VR than in R condition. The general sense of presence was reported as acceptable (mean value of 4.6 out of 6) for all the participants, and the cybersickness was reported as acceptable (SSQ mean value of 4.25 on the three circuits in VR condition). CONCLUSION: Driving performances were statistically different in the most complicated circuit C3 with an increased number of collisions in VR, but were not statistically different for the two easiest circuits C1 and C2 in R and VR conditions. In addition, there were no significant adverse effects such as cybersickness. The results show the value of the simulator for driving training applications. Still, the mental load was higher in VR than in R condition, thus mitigating the potential for use with people with cognitive disorders. Further studies should be conducted to assess the quality of skill transfer for novice drivers from the simulator to the real world. Trial registration Ethical approval n ∘ 2019-A001306-51 from Comité de Protection des Personnes Sud Mediterranée IV. Trial registered the 19/11/2019 on ClinicalTrials.gov in ID: NCT04171973.

Perceptual decoupling or trigger happiness: the effect of response delays and shorter presentation times on a go-no-go task with a high go prevalence.

In the current investigation, we modified the high Go, low No-Go Sustained Attention to Response Task (SART). Some researchers argue a commission error, an inappropriate response to a No-Go stimulus, in the SART is due to the participant being inattentive, or perceptually decoupled, during stimulus onset. Response delays in the SART reduce commission errors. A response delay may therefore enable a participant who is initially inattentive to recouple their attention in time to appropriately perceive the stimulus and withhold a response to a No-Go stimulus. However, shortening stimulus display duration in the SART should limit the possibility of the participant identifying the stimulus later, if they are initially not attending the stimulus. A response delay should not reduce commission errors if stimulus duration is kept to the minimum duration enabling stimulus recognition. In two experiments, we shortened stimulus onset to offset duration and added response delays of varying lengths. In both experiments, even when stimulus duration was shortened, response delays notably reduced commission errors if the delay was greater than 250 ms. In addition, using the Signal Detection Theory perspective in which errors of commission in the SART are due to a lenient response bias-trigger happiness, we predicted that response delays would result in a shift to a more conservative response bias in both experiments. These predictions were verified. The errors of commission in the SART may not be a measures of conscious awareness per se, but instead indicative of the level of participant trigger happiness-a lenient response bias.

Motor learning and performance in schizophrenia and aging: two different patterns of decline.

Psychomotor slowing has consistently been observed in schizophrenia, however research on motor learning in schizophrenia is limited. Additionally, motor learning in schizophrenia has never been compared with the waning of motor learning abilities in the elderly. Therefore, in an extensive study, 30 individuals with schizophrenia, 30 healthy age-matched controls and 30 elderly participants were compared on sensorimotor learning tasks including sequence learning and adaptation (both explicit and implicit), as well as tracking and aiming. This paper presents new findings on an explicit motor sequence learning task, an explicit verbal learning task and a simple aiming task and summarizes all previously published findings of this large investigation. Individuals with schizophrenia and elderly had slower Movement Time (MT)s compared with controls in all tasks, however both groups improved over time. Elderly participants learned slower on tracking and explicit sequence learning while individuals with schizophrenia adapted slower and to a lesser extent to movement perturbations in adaptation tasks and performed less well on cognitive tests including the verbal learning task. Results suggest that motor slowing is present in schizophrenia and the elderly, however both groups show significant but different motor skill learning. Cognitive deficits seem to interfere with motor learning and performance in schizophrenia while task complexity and decreased movement precision interferes with motor learning in the elderly, reflecting different underlying patterns of decline in these conditions. In addition, evidence for motor slowing together with impaired implicit adaptation supports the influence of cerebellum and the cerebello-thalamo-cortical-cerebellar (CTCC) circuits in schizophrenia, important for further understanding the pathophysiology of the disorder.

Movement-goal relevant object shape properties act as poor but viable cues for the attribution of motor errors to external objects.

When a context change is detected during motor learning, motor memories-internal models for executing movements within some context-may be created or existing motor memories may be activated and modified. Assigning credit to plausible causes of errors can allow for fast retrieval and activation of a motor memory, or a combination of motor memories, when the presence of such causes is detected. Features of the movement-context intrinsic to the movement dynamics, such as posture of the end effector, are often effective cues for detecting context change whereas features extrinsic to the movement dynamics, such as the colour of an object being moved, are often not. These extrinsic cues are typically not relevant to the motor task at hand and can be safely ignored by the motor system. We conducted two experiments testing if extrinsic but movement-goal relevant object-shape cues during an object-transport task can act as viable contextual cues for error assignment to the object, and the creation of new, object-shape-associated motor memories. In the first experiment we find that despite the object-shape cues, errors are primarily attributed to the hand transporting the object. In a second experiment, we find participants can execute differing movements cued by the object shape in a dual adaptation task, but the extent of adaptation is small, suggesting that movement-goal relevant object-shape properties are poor but viable cues for creating context specific motor memories.

Action initiation and action inhibition follow the same time course when compared under matched experimental conditions.

The ability to initiate an action quickly when needed and the ability to cancel an impending action are both fundamental to action control. It is often presumed that they are qualitatively distinct processes, yet they have largely been studied in isolation and little is known about how they relate to one another. Comparing previous experimental results shows a similar time course for response initiation and response inhibition. However, the exact time course varies widely depending on experimental conditions, including the frequency of different trial types and the urgency to respond. For example, in the stop-signal task, where both action initiation and action inhibition are involved and could be compared, action inhibition is typically found to be much faster. However, this apparent difference is likely due to there being much greater urgency to inhibit an action than to initiate one in order to avoid failing at the task. This asymmetry in the urgency between action initiation and action inhibition makes it impossible to compare their relative time courses in a single task. Here, we demonstrate that when action initiation and action inhibition are measured separately under conditions that are matched as closely as possible, their speeds are not distinguishable and are positively correlated across participants. Our results raise the possibility that action initiation and action inhibition may not necessarily be qualitatively distinct processes but may instead reflect complementary outcomes of a single decision process determining whether or not to act.NEW & NOTEWORTHY The time courses of initiating an action and canceling an action have largely been studied in isolation, and little is known about their relationship. Here, we show that when measured under comparable conditions the speeds of action initiation and action inhibition are the same. This finding raises the possibility that these two functions may be more closely related than previously assumed, with potentially important implications for their underlying neural basis.

Variable, sometimes absent, but never negative: Applying multilevel models of variability to the backward crosstalk effect to find theoretical constraints.

When performing two tasks at the same time, the congruency of the second task's features influences performance in the first task. This is called the backward crosstalk effect (BCE), a phenomenon that influences both theories of binding and of dual-task capacity limitations. The question of whether the BCE is found in all participants at all times is relevant for understanding the basis of the effect. For example, if the BCE is based on strategic choices, it can be variable, but if it is automatic and involuntary, it should never vary in whether it is present or not. Variability in observed BCE sizes was already documented and discussed when the group average effect was first reported (Hommel, 1998). Yet the theories discussed at the time did not motivate a more direct analysis of this variability, nor did the readily available statistical tools permit it. Some statistical approaches recently applied in cognitive psychology allow such a variability-focused analysis and some more recent theoretical debates would benefit from this as well. We assessed the variability of the BCE as well as a BCE-like free-choice congruency effect by applying a Bayesian multilevel modeling approach to the data from a dual-tasking experiment. Trials consisted of a two- and a four-response task. We manipulated which task was presented first and whether the response to the four-choice task was free or forced choice. RT data were best predicted by a model in which the BCE is zero in part of the population and drawn from a normal distribution truncated at zero (and thus always positive) in the rest of the population. Choice congruency bias data were best predicted by a model assuming this effect to be drawn from a normal distribution truncated at zero (but, in contrast to the RT data, without the subset of the population where it is zero). The BCE is not an inflexible and universal phenomenon that is directly linked to an inherent structural trait of human cognition. We discuss theoretical constraints implied by these results with a focus on what we can infer about the traits of the factors that influence BCE size. We suggest that future research might add further major constraints by using multi-session experiments to distinguish between-person and within-person variability. Our results show that the BCE is variable. The next step is understanding along which axes it is variable and why it varies.

Why is there an error negativity on correct trials? A reappraisal.

In healthy subjects, the Error Negativity (Ne) was initially reported on errors and on partial errors, only. Later on, application of the Laplacian transformation to EEG data unmasked a Ne-like wave (Nc) that shares a main generator with the Ne, suggesting that the Nc is just a small Ne. However, the reason why a small Ne would persist on correct responses remains unclear. Now, sometimes, subthreshold EMG activations in the muscles corresponding to correct responses (not strong enough to reach the response threshold) can precede full-blown correct responses. These "partially correct" activities seem to correspond to (force) execution errors, as they evoke a sizeable Ne. Within the frames of the Reward Value and Prediction Model or of the Predicted Response-Outcome model we propose that the action monitoring system evokes a Ne/Nc on correct responses because, even when a correct choice has been made, the accuracy of response (force) execution cannot be fully predicted. If this interpretation is correct, it can be assumed that, once these execution errors have been corrected, the correctness of the (full-blown) correcting response is highly predictable. Consequently, they should evoke a smaller Nc/Ne than "pure" correct responses. We show, that for the response thresholds set in the present experiment, the correcting response of the trials containing a partially correct activation evoke no identifiable Nc at all. Therefore it seems that there usually is an Error Negativity on correct trials because the correctness of response (force) execution cannot be fully predicted.

Longitudinal point-of-care assessment of psychomotor vigilance in children in the epilepsy monitoring unit.

The epilepsy monitoring unit (EMU) is a complex and dynamic operational environment, where the cognitive and behavioural consequences of medical and environmental changes often go unnoticed. The psychomotor vigilance task (PVT) has been used to detect changes in cognition and behaviour in numerous contexts, including among astronauts on spaceflight missions, pilots, and commercial drivers. Here, we piloted serial point-of-care administration of the PVT in children undergoing invasive monitoring in the EMU. Seven children completed the PVT throughout their hospital admission and their performance was associated with daily seizure counts, interictal epileptiform discharges, number of antiseizure medications (ASMs) administered, and sleep quality metrics. Using mixed-effects models, we found that PVT reaction time and accuracy were adversely affected by greater number of ASMs and interictal epileptiform activity. We show that serial point-of-care PVT is simple and feasible in the EMU and may enable greater understanding of individual patient responses to medical and environmental alterations, inform clinical decision-making, and support quality-improvement and research initiatives.

Convolutional neural networks reveal properties of reach-to-grasp encoding in posterior parietal cortex.

Deep neural networks (DNNs) are widely adopted to decode motor states from both non-invasively and invasively recorded neural signals, e.g., for realizing brain-computer interfaces. However, the neurophysiological interpretation of how DNNs make the decision based on the input neural activity is limitedly addressed, especially when applied to invasively recorded data. This reduces decoder reliability and transparency, and prevents the exploitation of decoders to better comprehend motor neural encoding. Here, we adopted an explainable artificial intelligence approach - based on a convolutional neural network and an explanation technique - to reveal spatial and temporal neural properties of reach-to-grasping from single-neuron recordings of the posterior parietal area V6A. The network was able to accurately decode 5 different grip types, and the explanation technique automatically identified the cells and temporal samples that most influenced the network prediction. Grip encoding in V6A neurons already started at movement preparation, peaking during movement execution. A difference was found within V6A: dorsal V6A neurons progressively encoded more for increasingly advanced grips, while ventral V6A neurons for increasingly rudimentary grips, with both subareas following a linear trend between the amount of grip encoding and the level of grip skills. By revealing the elements of the neural activity most relevant for each grip with no a priori assumptions, our approach supports and advances current knowledge about reach-to-grasp encoding in V6A, and it may represent a general tool able to investigate neural correlates of motor or cognitive tasks (e.g., attention and memory tasks) from single-neuron recordings.

Multiscale entropy in a 10-minute vigilance task.

Research has shown multiscale entropy, brain signal behavior across time scales, to reliably increase at lower time scales with time-on-task fatigue. However, multiscale entropy has not been examined in short vigilance tasks (i.e., ≤ 10 min). Addressing this gap, we examine multiscale entropy during a 10-minute Psychomotor Vigilance Test (PVT). Thirty-four participants provided neural data while completing the PVT. We compared the first 2 min of the task to the 7th and 8th minutes to avoid end-spurt effects. Results suggested increased multiscale entropy at lower time scales later compared to earlier in the task, suggesting multiscale entropy is a strong marker of time-on-task fatigue onset during short vigils. Separate analyses for Fast and Slow performers reveal differential entropy patterns, particularly over visual cortices. Here, observed brain-behavior linkage between entropy and reaction time for slow performers suggests that entropy assays over sensory cortices might have predictive value for fatigue onset or shifts from on- to off-task states.

Effects of a dual-task activity on gait parameters of people with and without intellectual disabilities.

BACKGROUND: The main objective of this study was to evaluate gait parameters in people with intellectual disability (ID) and without intellectual disability (WID) in two different walking conditions [single task vs. dual task (DT)]. A secondary aim was to evaluate the dual-task cost (DTC) that the DT causes in each group. METHODS: A total of 119 participants joined in this study: 56 ID (30 men) and 63 WID (30 men). The OptoGait system was used to assess gait. In addition, Witty photocells were added to assess gait under the DT condition. RESULTS: Single support time was lower for participants with ID (P < 0.01), while double support time was higher (P < 0.05). All coefficients of variation for gait parameters were higher in participants with ID. Additionally, changes in gait were observed in both groups during the DT condition compared with the single-task condition. These changes were larger for participants with ID in step length, double support time and gait speed (P < 0.001), resulting in a higher DTC in these variables in the ID group (P < 0.01). CONCLUSIONS: Both groups reduced gait performance in the DT condition. However, greater gait variability occurred in the ID group. In addition, DTC was higher for the ID group in all variables analysed. Therefore, people with ID show worse gait performance during a DT than people WID.

Value Estimation versus Effort Mobilization: A General Dissociation between Ventromedial and Dorsomedial Prefrontal Cortex.

Deciding on a course of action requires both an accurate estimation of option values and the right amount of effort invested in deliberation to reach sufficient confidence in the final choice. In a previous study, we have provided evidence, across a series of judgment and choice tasks, for a dissociation between the ventromedial prefrontal cortex (vmPFC), which would represent option values, and the dorsomedial prefrontal cortex (dmPFC), which would represent the duration of deliberation. Here, we first replicate this dissociation and extend it to the case of an instrumental learning task, in which 24 human volunteers (13 women) choose between options associated with probabilistic gains and losses. According to fMRI data recorded during decision-making, vmPFC activity reflects the sum of option values generated by a reinforcement learning model and dmPFC activity the deliberation time. To further generalize the role of the dmPFC in mobilizing effort, we then analyze fMRI data recorded in the same participants while they prepare to perform motor and cognitive tasks (squeezing a handgrip or making numerical comparisons) to maximize gains or minimize losses. In both cases, dmPFC activity is associated with the output of an effort regulation model, and not with response time. Taken together, these results strengthen a general theory of behavioral control that implicates the vmPFC in the estimation of option values and the dmPFC in the energization of relevant motor and cognitive processes.

The effect of quiet eye training on golf putting performance in pressure situation.

To explores the effect and mechanism of quiet eye training on the accuracy of golfers´ putts in pressure situations and provides methods and basis for targeted attention training and control. 22 young golfers in China golf team aged from 13 to 18 were randomly assigned to the experimental group (quiet eye training group) and the control group (technical guidance group) according to gender. Both groups of participants underwent two consecutive weeks of push training (3 sets per day, 20 golf putts per set, rest for 3 min between sets) separately in accordance with the guidance of a professional psychological research group and an expert coach. Eye tracking technology, biofeedback technology, and subjective evaluation methods were used to test and analyze the push process of the two groups of participants before and after training under pressure situations (Eye movement behaviors and the heart rate were recorded by ASL Mobile Eye-XG and NeXus-2 biofeedback, pressure and state anxiety were evaluated by self-rating pressure scale and S-AI. Golf putting performance was recorded by a research graduate assistant). A higher hit ratio as well as lower pressure and SAI level was founded in quiet eye training group in the pressure situation, the quiet eye movement time and total fixation time was longer than technical group. The quiet eye training group has a better putting performance. Quiet eye training can improve the golf putting performance in pressure situations. After quiet eye training, the state anxiety decreased, the quiet eye movement time and the total fixation time increased in pressure situations.

Proximity to rewards modulates parameters of effortful control exertion.

The now-classic goal-gradient hypothesis posits that organisms increase effort expenditure as a function of their proximity to a goal. Despite nearly a century having passed since its original formulation, goal-gradient-like behavior in human cognitive performance remains poorly understood: Are we more willing to engage in costly cognitive processing when we are near, versus far, from a goal state? Moreover, the computational mechanisms underpinning these potential goal-gradient effects-for example, whether goal proximity affects fidelity of stimulus encoding, response caution, or other identifiable mechanisms governing speed and accuracy-are unclear. Here, in two experiments, we examine the effect of goal proximity, operationalized as progress toward the completion of a rewarded task block, upon task performance in an attentionally demanding oddball task. Supporting the goal-gradient hypothesis, we found that participants responded more quickly, but not less accurately, when rewards were proximal than when they were distal. Critically, this effect was only observed when participants were given information about goal proximity. Using hierarchical drift diffusion modeling, we found that these apparent goal-gradient performance effects were best explained by a collapsing bound model, in which proximity to a goal reduced response caution and increased information processing. Taken together, these results suggest that goal gradients could help explain the oft-observed fluctuations in engagement of cognitively effortful processing, extending the scope of the goal-gradient hypothesis to the domain of cognitive tasks. (PsycInfo Database Record (c) 2024 APA, all rights reserved).

Temporal recalibration in response to delayed visual feedback of active versus passive actions: an fMRI study.

The brain can adapt its expectations about the relative timing of actions and their sensory outcomes in a process known as temporal recalibration. This might occur as the recalibration of timing between the sensory (e.g. visual) outcome and (1) the motor act (sensorimotor) or (2) tactile/proprioceptive information (inter-sensory). This fMRI recalibration study investigated sensorimotor contributions to temporal recalibration by comparing active and passive conditions. Subjects were repeatedly exposed to delayed (150 ms) or undelayed visual stimuli, triggered by active or passive button presses. Recalibration effects were tested in delay detection tasks, including visual and auditory outcomes. We showed that both modalities were affected by visual recalibration. However, an active advantage was observed only in visual conditions. Recalibration was generally associated with the left cerebellum (lobules IV, V and vermis) while action related activation (active > passive) occurred in the right middle/superior frontal gyri during adaptation and test phases. Recalibration transfer from vision to audition was related to action specific activations in the cingulate cortex, the angular gyrus and left inferior frontal gyrus. Our data provide new insights in sensorimotor contributions to temporal recalibration via the middle/superior frontal gyri and inter-sensory contributions mediated by the cerebellum.

Flanker interference at both stimulus and response levels decreases with age.

When trying to identify the colour of a target, people's performance is impaired by nearby distractors of different colours. It is controversial whether these interference effects originate from competing stimuli, competing responses or from both simultaneously. These interference effects may also differ depending on a person's age. Comparisons between studies show mixed results, while differences in experimental design and data analysis complicate the interpretation. In our study, we manipulated the relative proportions of congruent and incongruent trials with respect to both stimuli and responses. Considering this aspect, we asked whether people resolve stimulus and response interference differently at different ages. 92 children (6-14 years), 25 young adults (20-43 years) and 33 older adults (60-84 years) performed a coloured version of the Eriksen flanker task. Since reaction times and errors were correlated, inverse efficiency scores were used to address speed-accuracy trade-offs between groups. Absolute interference effects were used to measure relationships with age. The results showed first, unexpectedly, that response interference was comparable between stimulus- and response-balanced conditions. Second, performance at all ages was significantly influenced both by competing stimuli and responses. Most importantly, the size of interference effects decreased with age. These findings cast some doubt on the conclusions of previous studies, and raise further questions about how cognitive control is best measured across the lifespan.