Predicting the metabolic energy costs of bipedalism using evolutionary robotics

To understand the evolution of bipedalism among the homnoids in an ecological context we need to be able to estimate theenerrgetic cost of locomotion in fossil forms. Ideally such an estimate would be based entirely on morphology since, except for the rare instances where footprints are preserved, this is hte only primary source of evidence available. In this paper we use evolutionary robotics techniques (genetic algoritms, pattern generators and mechanical modeling) to produce a biomimentic simulation of bipedalism based on human body dimensions. The mechnaical simulation is a seven-segment, two-dimensional model with motive force provided by tension generators representing the major muscle groups acting around the lower-limb joints. Metabolic energy costs are calculated from the muscel model, and bipedal gait is generated using a finite-state pattern generator whose parameters are produced using a genetic algorithm with locomotor economy (maximum distance for a fixed energy cost) as the fitness criterion. The model is validated by comparing the values it generates with those for modern humans. The result (maximum efficiency of 200 J m-1) is within 15% of the experimentally derived value, which is very encouraging and suggests that this is a useful analytic technique for investigating the locomotor behaviour of fossil forms. Initial work suggests that in the future this technique could be used to estimate other locomotor parameters such as top speed. In addition, the animations produced by this technique are qualitatively very convincing, which suggests that this may also be a useful technique for visualizing bipedal locomotion.

The effect of knee flexion contracture following total knee arthroplasty on the energy cost of walking.

This study evaluated the energy cost of walking (Cw) with knee flexion contractures (FC) simulated with a knee brace, in total knee arthroplasty (TKA) recipients (n=16) and normal controls (n=15), and compared it to baseline (no brace). There was no significant difference in Cw between the groups at baseline but TKA recipients walked slower (P=0.048) and with greater knee flexion in this condition (P=0.003). Simulated FC significantly increased Cw in both groups (TKA P=0.020, control P=0.002) and this occurred when FC exceeded 20° in the TKA group and 15° in the controls. Reported perceived exertion was only significantly increased by FC in the control group (control P<0.001, TKA P=0.058). Simulated knee FCs less than 20° do not increase Cw or perceived exertion in TKA recipients.

Association between energy cost of walking, muscle activation, and biomechanical parameters in older female fallers and non-fallers

Objective: To determine the nervous activation, muscle strength, and biomechanical parameters that influence the cost of walking in older fallers and non-fallers. Methods: Maximal voluntary isokinetic torque was measured for the hip, knee and ankle of older women. Oxygen consumption was measured at rest and during 8 min of walking at self-selected speed. An additional minute of walking was performed to collect kinematic variables and the electromyographic signal of trunk, hip, knee, and ankle muscles, which was analyzed by the linear envelope. Cost of walking was calculated by subtracting resting body mass-normalized oxygen consumption from walking body mass-normalized oxygen consumption. Stride time and length, and ankle and hip range of motion were calculated from kinematic data. Findings: Older adult fallers had 28% lower knee extensor strength (p = 0.02), 47% lower internal oblique activation at heel contact (p = 0.03), and higher coactivation between tibialis anterior and gastrocnemius lateralis in each of the gait phases (p < 0.05). For fallers, a higher activation of gluteus maximus was associated with a higher cost of walking (r = 0.55, p < 0.05 and r = 0.71, p < 0.01, before and after heel contact, respectively). For non-fallers, an association between cost of walking and age (r = 0.60, p = 0.01) and cost of walking and thigh muscle coactivation (r = 0.53, p = 0.01) existed. Interpretation: This study demonstrated that there may be links between lower-extremity muscle weakness, muscle activation patterns, altered gait, and increased cost of walking in older fallers. © 2013 Elsevier Ltd. All rights reserved.

Excess body weight and gait influence energy cost of walking in older adults

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Relations entre la mobilité, la condition physique et les fonctions cognitives dans une perspective de vieillissement sain

Thèse effectuée en cotutelle (Université de Poitiers, France)

The energy costs of sexual dimorphism in mole-rats are morphological not behavioural.

Different reproductive strategies of males and females may lead to the evolution of differences in their energetic costs of reproduction, overall energetic requirements and physiological performances. Sexual dimorphism is often associated with costly behaviours (e.g. large males might have a competitive advantage in fighting, which is energetically expensive). However, few studies of mammals have directly compared the energy costs of reproductive activities between sexes. We compared the daily energy expenditure (DEE) and resting metabolic rate (RMR) of males and females of two species of mole-rat, Bathyergus janetta and Georychus capensis (the former is sexually dimorphic in body size and the latter is not) during a period of intense digging when males seek females. We hypothesized that large body size might be indicative of greater digging or fighting capabilities, and hence greater mass-independent DEE values in males of the sexually dimorphic species. In contrast to this prediction, although absolute values of DEE were greater in B. janetta males, mass-independent values were not. No differences were apparent between sexes in G. capensis. By comparison, although RMR values were greater in B. janetta than G. capensis, no differences were apparent between the sexes for either species. The energy cost of dimorphism is most likely to be the cost of maintenance of a large body size, and not the cost of behaviours performed when an individual is large.

Experimental ocean acidification alters the allocation of metabolic energy

Energy is required to maintain physiological homeostasis in response to environmental change. Although responses to environmental stressors frequently are assumed to involve high metabolic costs, the biochemical bases of actual energy demands are rarely quantified. We studied the impact of a near-future scenario of ocean acidification [800 µatm partial pressure of CO2 (pCO2)] during the development and growth of an important model organism in developmental and environmental biology, the sea urchin Strongylocentrotus purpuratus. Size, metabolic rate, biochemical content, and gene expression were not different in larvae growing under control and seawater acidification treatments. Measurements limited to those levels of biological analysis did not reveal the biochemical mechanisms of response to ocean acidification that occurred at the cellular level. In vivo rates of protein synthesis and ion transport increased 50% under acidification. Importantly, the in vivo physiological increases in ion transport were not predicted from total enzyme activity or gene expression. Under acidification, the increased rates of protein synthesis and ion transport that were sustained in growing larvae collectively accounted for the majority of available ATP (84%). In contrast, embryos and prefeeding and unfed larvae in control treatments allocated on average only 40% of ATP to these same two processes. Understanding the biochemical strategies for accommodating increases in metabolic energy demand and their biological limitations can serve as a quantitative basis for assessing sublethal effects of global change. Variation in the ability to allocate ATP differentially among essential functions may be a key basis of resilience to ocean acidification and other compounding environmental stressors.

Measurement of energy expenditure of daily tasks among mothers of young children

There is currently some debate about whether the energy expenditure of domestic tasks is sufficient to confer health benefits. The aim of this study was therefore to measure the energy cost of five activities commonly undertaken by mothers of young children. Seven women with at least one child younger than five years of age spent 15 minutes in each of the following activities: sitting quietly, vacuum cleaning, washing windows, walking at moderate pace (approx 5km/hour), walking with a stroller and grocery shopping in a super-market. Each of the six 'trials' was completed on the same day, in random order. A carefully calibrated portable gas analyser was used to measure oxygen uptake during each activity, and data were converted to units of energy expenditure (METS). Vacuum cleaning, washing windows and walking with and without a stroller were found to be 'moderate intensity activities' (3 to 6 METs), but supermarket shopping did not reach this criterion. The MET values for these activities were similar to those reported in the Compendium of Physical Activities (Ainsworth et al., 2000). However, the energy expenditures of walking, both with and without a stroller, were higher than those reported in the Compendium. The findings suggest that some of the tasks associated with domestic caring duties are conducted at an intensity which is sufficient to confer some health benefit. Such benefits will only accrue however if the daily duration of these activities is sufficient to meet current guidelines.

Objective analysis of gait coordination for total knee arthroplasty outcome

Introduction: Coordination is a strategy chosen by the central nervous system to control the movements and maintain stability during gait. Coordinated multi-joint movements require a complex interaction between nervous outputs, biomechanical constraints, and pro-prioception. Quantitatively understanding and modeling gait coordination still remain a challenge. Surgeons lack a way to model and appreciate the coordination of patients before and after surgery of the lower limbs. Patients alter their gait patterns and their kinematic synergies when they walk faster or slower than normal speed to maintain their stability and minimize the energy cost of locomotion. The goal of this study was to provide a dynamical system approach to quantitatively describe human gait coordination and apply it to patients before and after total knee arthroplasty. Methods: A new method of quantitative analysis of interjoint coordination during gait was designed, providing a general model to capture the whole dynamics and showing the kinematic synergies at various walking speeds. The proposed model imposed a relationship among lower limb joint angles (hips and knees) to parameterize the dynamics of locomotion of each individual. An integration of different analysis tools such as Harmonic analysis, Principal Component Analysis, and Artificial Neural Network helped overcome high-dimensionality, temporal dependence, and non-linear relationships of the gait patterns. Ten patients were studied using an ambulatory gait device (Physilog®). Each participant was asked to perform two walking trials of 30m long at 3 different speeds and to complete an EQ-5D questionnaire, a WOMAC and Knee Society Score. Lower limbs rotations were measured by four miniature angular rate sensors mounted respectively, on each shank and thigh. The outcomes of the eight patients undergoing total knee arthroplasty, recorded pre-operatively and post-operatively at 6 weeks, 3 months, 6 months and 1 year were compared to 2 age-matched healthy subjects. Results: The new method provided coordination scores at various walking speeds, ranged between 0 and 10. It determined the overall coordination of the lower limbs as well as the contribution of each joint to the total coordination. The difference between the pre-operative and post-operative coordination values were correlated with the improvements of the subjective outcome scores. Although the study group was small, the results showed a new way to objectively quantify gait coordination of patients undergoing total knee arthroplasty, using only portable body-fixed sensors. Conclusion: A new method for objective gait coordination analysis has been developed with very encouraging results regarding the objective outcome of lower limb surgery.

Fatores intrínsecos do custo energético da locomoção durante a natação

The amount of metabolic energy spent in transporting the body mass of the subject over a unit of distance has been defined as the energy cost of locomotion, or regarding to swimming, cost of swimming. The differences in the cost of swimming between the individuals seem to be influenced by two main factors, the hydrodynamic resistance and technical skill of the swimmer. The lower cost of swimming showed by females has been attributed to a smaller hydrodynamic resistance due to their smaller size, larger percentage fat and more streamlined position. However, the difference in cost of swimming between males and females disappears when correcting for body size. With regard to children, the higher energy cost of swimming when correcting for body size may be caused by the lower swimming technique showed by them. For individuals with the same anthropometric characteristics, the better swimming technique and larger size of propelling surface, associated with higher propelling efficiency, may decrease the energy cost of swimming. When comparing different types of strokes, the most economical stroke is crawl, followed by backstroke, irrespective the swimming velocity. Butterfly is the less economical at low velocities (< 0.8 m·s-1). However, above that velocity the breaststroke become the less economical stroke.

Running behavior and its energy cost in mice selectively bred for high voluntary locomotor activity

Locomotion is central to behavior and intrinsic to many fitnesscritical activities (e.g., migration, foraging), and it competes with other life-history components for energy. However, detailed analyses of how changes in locomotor activity and running behavior affect energy budgets are scarce. We quantified these effects in four replicate lines of house mice that have been selectively bred for high voluntary wheel running (S lines) and in their four nonselected control lines (C lines). We monitored wheel speeds and oxygen consumption for 24-48 h to determine daily energy expenditure (DEE), resting metabolic rate (RMR), locomotor costs, and running behavior (bout characteristics). Daily running distances increased roughly 50%-90% in S lines in response to selection. After we controlled for body mass effects, selection resulted in a 23% increase in DEE in males and a 6% increase in females. Total activity costs (DEE - RMR) accounted for 50%-60% of DEE in both S and C lines and were 29% higher in S males and 5% higher in S females compared with their C counterparts. Energetic costs of increased daily running distances differed between sexes because S females evolved higher running distances by running faster with little change in time spent running, while S males also spent 40% more time running than C males. This increase in time spent running impinged on high energy costs because the majority of running costs stemmed from postural costs (the difference between RMR and the zero-speed intercept of the speed vs. metabolic rate relationship). No statistical differences in these traits were detected between S and C females, suggesting that large changes in locomotor behavior do not necessarily effect overall energy budgets. Running behavior also differed between sexes: within S lines, males ran with more but shorter bouts than females. Our results indicate that selection effects on energy budgets can differ dramatically between sexes and that energetic constraints in S males might partly explain the apparent selection limit for wheel running observed for over 15 generations. © 2009 by The University of Chicago. All rights reserved.

Interactions between energy intake and expenditure in the development and treatment of obesity.

Summary There are four interactions to consider between energy intake (EI) and energy expenditure (EE) in the development and treatment of obesity. (1) Does sedentariness alter levels of EI or subsequent EE? and (2) Do high levels of EI alter physical activity or exercise? (3) Do exercise-induced increases in EE drive EI upwards and undermine dietary approaches to weight management and (4) Do low levels of EI elevate or decrease EE? There is little evidence that sedentariness alters levels of EI. This lack of cross-talk between altered EE and EI appears to promote a positive EB. Lifestyle studies also suggest that a sedentary routine actually offers the opportunity for over-consumption. Substantive changes in non exercise activity thermogenesis are feasible, but not clearly demonstrated. Cross talk between elevated EE and EI is initially too weak and takes too long to activate, to seriously threaten dietary approaches to weight management. It appears that substantial fat loss is possible before intake begins to track a sustained elevation of EE. There is more evidence that low levels of EI does lower physical activity levels, in relatively lean men under conditions of acute or prolonged semi-starvation and in dieting obese subjects. During altered EB there are a number of small but significant changes in the components of EE, including (i) sleeping and basal metabolic rate, (ii) energy cost of weight change alters as weight is gained or lost, (iii) exercise efficiency, (iv) energy cost of weight bearing activities, (v) during substantive overfeeding diet composition (fat versus carbohydrate) will influence the energy cost of nutrient storage by ~ 15%. The responses (i-v) above are all “obligatory” responses. Altered EB can also stimulate facultative behavioural responses, as a consequence of cross-talk between EI and EE. Altered EB will lead to changes in the mode duration and intensity of physical activities. Feeding behaviour can also change. The degree of inter-individual variability in these responses will define the scope within which various mechanisms of EB compensation can operate. The relative importance of “obligatory” versus facultative, behavioural responses -as components of EB control- need to be defined.

Minimum cost of transport in human running is not ubiquitous

Purpose This study explores recent claims that humans exhibit a minimum cost of transport (CoTmin) for running which occurs at an intermediate speed, and assesses individual physiological, gait and training characteristics. Methods Twelve healthy participants with varying levels of fitness and running experience ran on a treadmill at six self-selected speeds in a discontinuous protocol over three sessions. Running speed (km[middle dot]hr-1), V[spacing dot above]O2 (mL[middle dot]kg-1[middle dot]km-1), CoT (kcal[middle dot]km-1), heart rate (beats[middle dot]min-1) and cadence (steps[middle dot]min-1) were continuously measured. V[spacing dot above]O2 max was measured on a fourth testing session. The occurrence of a CoTmin was investigated and its presence or absence examined with respect to fitness, gait and training characteristics. Results Five participants showed a clear CoTmin at an intermediate speed and a statistically significant (p < 0.05) quadratic CoT-speed function, while the other participants did not show such evidence. Participants were then categorized and compared with respect to the strength of evidence for a CoTmin (ClearCoTmin and NoCoTmin). The ClearCoTmin group displayed significantly higher correlation between speed and cadence; more endurance training and exercise sessions per week; than the NoCoTmin group; and a marginally non-significant but higher aerobic capacity. Some runners still showed a CoTmin at an intermediate speed even after subtraction of resting energy expenditure. Conclusion The findings confirm the existence of an optimal speed for human running, in some but not all participants. Those exhibiting a COTmin undertook a higher volume of running, ran with a cadence that was more consistently modulated with speed, and tended to be aerobically fitter. The ability to minimise the energetic cost of transport appears not to be ubiquitous feature of human running but may emerge in some individuals with extensive running experience.