It is not just the work you do, but how you do it: the metabolic cost of walking uphill and downhill with varying grades.

The cost of walking and running on uneven terrain is not directly explained by external mechanical work. Although metabolic cost of transport increases linearly with gradient at uphill and downhill gradients exceeding 15%, at shallower gradients, the relationship is nonlinear, with the minimum cost occurring at ∼10% downhill grade. Given these nonlinear relationships between grade and metabolic cost, we projected a significant difference in the total metabolic cost of two walking conditions that required the same total external mechanical work be performed over the same total period of time; in one condition, time was spent walking to gradients that were fixed at +10.5% and -10.5% and in the other condition time was spent walking to gradients that varied from 0 to +21% and from -21 to 0%. We compared these two conditions experimentally, using an approach to quantify nonsteady-state oxidative energy expenditure. In line with our projection, the "variable" grade condition resulted in an 8.3 ± 2.2% higher total cumulative oxidative energy expenditure (J·kg-1) compared with the "fixed" grade condition (P < 0.001). Future work should aim to apply our approach across different gradients, speeds, and forms of locomotion; especially those that might provide insight into how humans optimize locomotion on variable grade routes.NEW & NOTEWORTHY We use a method for quantifying nonsteady-state energetics to show that regardless of whether the same total gain and loss in elevation (i.e., same total external mechanical work) is achieved over the same period of time, the total energy expenditure of different graded walking conditions can vary depending on the grades that are walked at and for how long they are walked at.

Validation of a musculoskeletal model for simulating muscle mechanics and energetics during diverse human hopping tasks.

Computational musculoskeletal modelling has emerged as an alternative, less-constrained technique to indirect calorimetry for estimating energy expenditure. However, predictions from modelling tools depend on many assumptions around muscle architecture and function and motor control. Therefore, these tools need to continue to be validated if we are to eventually develop subject-specific simulations that can accurately and reliably model rates of energy consumption for any given task. In this study, we used OpenSim software and experimental motion capture data to simulate muscle activations, muscle fascicle dynamics and whole-body metabolic power across mechanically and energetically disparate hopping tasks, and then evaluated these outputs at a group- and individual-level against experimental electromyography, ultrasound and indirect calorimetry data. Comparing simulated and experimental outcomes, we found weak to strong correlations for peak muscle activations, moderate to strong correlations for absolute fascicle shortening and mean shortening velocity, and strong correlations for gross metabolic power. These correlations tended to be stronger on a group-level rather than individual-level. We encourage the community to use our publicly available dataset from SimTK.org to experiment with different musculoskeletal models, muscle models, metabolic cost models, optimal control policies, modelling tools and algorithms, data filtering etc. with subject-specific simulations being a focal goal.

Linking muscle mechanics to the metabolic cost of human hopping.

Many models have been developed to predict metabolic energy expenditure based on biomechanical proxies of muscle function. However, current models may only perform well for select forms of locomotion, not only because the models are rarely rigorously tested across subtle and broad changes in locomotor task but also because previous research has not adequately characterised different forms of locomotion to account for the potential variability in muscle function and thus metabolic energy expenditure. To help to address the latter point, the present study imposed frequency and height constraints to hopping and quantified gross metabolic power as well as the activation requirements of medial gastrocnemius (MG), lateral gastrocnemius (GL), soleus (SOL), tibialis anterior (TA), vastus lateralis (VL), rectus femoris (RF) and biceps femoris (BF), and the work requirements of GL, SOL and VL. Gross metabolic power increased with a decrease in hop frequency and increase in hop height. There was no hop frequency or hop height effect on the mean electromyography (EMG) data of ankle musculature; however, the mean EMG of VL and RF increased with a decrease in hop frequency and that of BF increased with an increase in hop height. With a reduction in hop frequency, GL, SOL and VL fascicle shortening, fascicle shortening velocity and fascicle to MTU shortening ratio increased, whereas with an increase in hop height, only SOL fascicle shortening velocity increased. Therefore, within the constraints that we imposed, decreases in hop frequency and increases in hop height resulted in increases in metabolic power that could be explained by increases in the activation requirements of knee musculature and/or increases in the work requirements of both knee and ankle musculature.