Slowing of Movements in Healthy Aging as a Rational Economic Response to an Elevated Effort Landscape.

Why do we move slower as we grow older? The reward circuits of the brain, which tend to invigorate movements, decline with aging, raising the possibility that reduced vigor is due to the diminishing value that our brain assigns to movements. However, as we grow older, it also becomes more effortful to make movements. Is age-related slowing principally a consequence of increased effort costs from the muscles, or reduced valuation of reward by the brain? Here, we first quantified the cost of reaching via metabolic energy expenditure in human participants (male and female), and found that older adults consumed more energy than the young at a given speed. Thus, movements are objectively more costly for older adults. Next, we observed that when reward increased, older adults, like the young, responded by initiating their movements earlier. Yet, unlike the young, they were unwilling to increase their movement speed. Was their reluctance to reach quicker for rewards due to the increased effort costs, or because they ascribed less value to the movement? Motivated by a mathematical model, we next made the young experience a component of aging by making their movements more effortful. Now the young responded to reward by reacting faster but chose not to increase their movement speed. This suggests that slower movements in older adults are partly driven by an adaptive response to an elevated effort landscape. Moving slower may be a rational economic response the brain is making to mitigate the elevated effort costs that accompany aging.

Long-term outcomes in patients treated with tissue-sparing posterior cervical fusion to revise a 1-level pseudarthrosis following ACDF.

STUDY DESIGN: Observational Study BACKGROUND: Symptomatic pseudarthrosis is one long-term complication in patients treated with anterior discectomy and fusion (ACDF). When revising a pseudarthrosis, a surgeon must decide to intervene posteriorly and/or anteriorly. Open posterior cervical fusion (PCF) is attractive for high rates of arthrodesis, however this technique introduces risks of added complications resulting from extensive soft tissue dissection. The purpose of this study was to assess long-term outcomes in patients undergoing tissue-sparing PCF with facet instrumentation to treat a single level pseudarthrosis. METHODS: Forty-five subjects were recruited from six participating sites. All subjects had a history of ACDF that was subsequently revised with tissue-sparing PCF to treat symptomatic pseudarthrosis at one level. Long-term radiographic assessments included flexion and extension X-ray and multi-planar CT. Subjects additionally completed a patient satisfaction questionnaire. Radiographs were assessed by investigators and an independent core imaging lab to diagnose implant integrity and arthrodesis at the revised levels. RESULTS: The revision procedure required a median 49 min to complete with an estimated blood loss of 10 cc. Subjects were discharged a median 1 day following treatment. There were no instances of hospital re-admission nor subsequent surgical interventions. Study follow-up assessments were performed a median 39 months from revision. Surgeons diagnosed complete fusion in 91 % of cases. The core imaging lab identified bridging bone across the revised segment in 80 % of cases. Range of motion was < 2° in 93 % of cases. Seventy-four percent of subjects reported being satisfied with their outcomes. CONCLUSIONS: This study summarizes long-term radiographic outcomes in a cohort of patients receiving tissue-sparing PCF for the treatment of pseudarthrosis. Assessed years after revision, patients achieved rates of arthrodesis similar to open PCF without the soft tissue dissection responsible for perioperative morbidity and long-term soft tissue pain.

Effort cost of reaching prompts vigor reduction in older adults.

As people age, they move slower. Is age-related reduction in vigor a reflection of a reduced valuation of reward by the brain, or a consequence of increased effort costs by the muscles? Here, we quantified cost of movements objectively via the metabolic energy that young and old participants consumed during reaching and found that in order reach at a given speed, older adults expended more energy than the young. We next quantified how reward modulated movements in the same populations and found that like the young, older adults responded to increased reward by initiating their movements earlier. Yet, their movements were less sensitive to increased reward, resulting in little or no modulation of reach speed. Lastly, we quantified the effect of increased effort on how reward modulated movements in young adults. Like the effects of aging, when faced with increased effort the young adults responded to reward primarily by reacting faster, with little change in movement speed. Therefore, reaching required greater energetic expenditure in the elderly, suggesting that the slower movements and reactions exhibited in aging are partly driven by an adaptive response to an elevation in the energetic landscape of effort. That is, moving slower appears to be a rational economic consequence of aging. Significance statement: Healthy aging coincides with a reduction in speed, or vigor, of walking, reaching, and eye movements. Here we focused on disentangling two opposing sources of aging-related movement slowing: reduced reward sensitivity due to loss of dopaminergic tone, or increased energy expenditure movements related to mitochondrial or muscular inefficiencies. Through a series of three experiments and construction of a computational model, here we demonstrate that transient changes in reaction time and movement speed together offer a quantifiable metric to differentiate between reward- and effort-based alterations in movement vigor. Further, we suggest that objective increases in the metabolic cost of moving, not reductions in reward valuation, are driving much of the movement slowing occurring alongside healthy aging.

Using metabolic energy to quantify the subjective value of physical effort.

Economists have known for centuries that to understand an individual's decisions, we must consider not only the objective value of the goal at stake, but its subjective value as well. However, achieving that goal ultimately requires expenditure of effort. Surprisingly, despite the ubiquitous role of effort in decision-making and movement, we currently do not understand how effort is subjectively valued in daily movements. Part of the difficulty arises from the lack of an objective measure of effort. Here, we use a physiological approach to address this knowledge gap. We quantified objective effort costs by measuring metabolic cost via expired gas analysis as participants performed a reaching task against increasing resistance. We then used neuroeconomic methods to quantify each individual's subjective valuation of effort. Rather than the diminishing sensitivity observed in reward valuation, effort was valued objectively, on average. This is significantly less than the near-quadratic sensitivity to effort observed previously in force-based motor tasks. Moreover, there was significant inter-individual variability with many participants undervaluing or overvaluing effort. These findings demonstrate that in contrast with monetary decisions in which subjective value exhibits diminishing marginal returns, effort costs are valued more objectively in low-effort reaching movements common in daily life.

Movement Vigor as a Reflection of Subjective Economic Utility.

To understand subjective evaluation of an option, various disciplines have quantified the interaction between reward and effort during decision making, producing an estimate of economic utility, namely the subjective 'goodness' of an option. However, variables that affect utility of an option also influence the vigor of movements toward that option. For example, expectation of reward increases speed of saccadic eye movements, whereas expectation of effort decreases this speed. These results imply that vigor may serve as a new, real-time metric with which to quantify subjective utility, and that the control of movements may be an implicit reflection of the brain's economic evaluation of the expected outcome.

Contributions of metabolic and temporal costs to human gait selection.

Humans naturally select several parameters within a gait that correspond with minimizing metabolic cost. Much less is understood about the role of metabolic cost in selecting between gaits. Here, we asked participants to decide between walking or running out and back to different gait specific markers. The distance of the walking marker was adjusted after each decision to identify relative distances where individuals switched gait preferences. We found that neither minimizing solely metabolic energy nor minimizing solely movement time could predict how the group decided between gaits. Of our twenty participants, six behaved in a way that tended towards minimizing metabolic energy, while eight favoured strategies that tended more towards minimizing movement time. The remaining six participants could not be explained by minimizing a single cost. We provide evidence that humans consider not just a single movement cost, but instead a weighted combination of these conflicting costs with their relative contributions varying across participants. Individuals who placed a higher relative value on time ran faster than individuals who placed a higher relative value on metabolic energy. Sensitivity to temporal costs also explained variability in an individual's preferred velocity as a function of increasing running distance. Interestingly, these differences in velocity both within and across participants were absent in walking, possibly due to a steeper metabolic cost of transport curve. We conclude that metabolic cost plays an essential, but not exclusive role in gait decisions.

Vigor of reaching movements: reward discounts the cost of effort.

Making a movement may be thought of as an economic decision in which one spends effort to acquire reward. Time discounts reward, which predicts that the magnitude of reward should affect movement vigor: we should move faster, spending greater effort, when there is greater reward at stake. Indeed, saccade peak velocities are greater and reaction-times shorter when a target is paired with reward. In this study, we focused on human reaching and asked whether movement kinematics were affected by expectation of reward. Participants made out-and-back reaching movements to one of four quadrants of a 14-cm circle. During various periods of the experiment only one of the four quadrants was paired with reward, and the transition from reward to nonreward status of a quadrant occurred randomly. Our experiment design minimized dependence of reward on accuracy, granting the subjects wide latitude in self-selecting their movement speed, amplitude, and variability. When a quadrant was paired with reward, reaching movements had a shorter reaction time, higher peak velocity, and greater amplitude. Despite this greater vigor, movements toward the rewarded quadrant suffered from less variability: both reaction times and reach kinematics were less variable when there was expectation of reward. Importantly, the effect of reward on vigor was specific to the movement component that preceded the time of reward (outward reach), not the movement component that followed it (return reach). Our results suggest that expectation of reward not only increases vigor of human reaching but also decreases its variability. NEW & NOTEWORTHY Movements may be thought of as an economic transaction where the vigor of the movement represents the effort that the brain is willing to expend to acquire a rewarding state. We show that in reaching, reward discounts the cost of effort, producing movements with shorter reaction time, higher velocity, greater amplitude, and reduced reaction-time variability. These results complement earlier observations in saccades, suggesting a common principle of economics across modalities of motor control.