The relation between maximal running speed and body mass in terrestrial mammals

The available data on maximal running speeds of mammals are presented, and the relationship between speed and body mass is considered. For all mammals ( n = 106), maximal running speed scales as (body mass)^0–17; however, the largest mammals are not the fastest, and an optimal size with regards to running ability is suggested ( 119 kg). Maximal running speeds are, on the average, somewhat more than twice maximal aerobic speeds.
Within the Artiodactyla, Carnivora or Rodentia, maximal running speed is mass independent, in agreement with theoretical expectations for geometrically similar animals (Thompson, 1917; Hill, 1950). McMahon's (1975 b ) model for elastic similarity is therefore not supported by the available data on maximal running speeds, and there appears to be no necessary correspondence between scaling of limb bone proportions and running ability.     

跑台运动促进幼龄大鼠学习能力

为了探讨跑台运动对幼龄大鼠学习能力的影响,实验采用5周龄Sprague-Dawley大鼠,随机分为安静对照组和跑台运动组,其中跑台运动组大鼠以低强度进行为期一周的跑台运动;然后使用Morris水迷宫,对两组大鼠的定位航行和空间探索能力进行分析。在定位航行实验中,运动组大鼠寻找到平台的潜伏期明显短于对照组（P〈0．05）;并且随着训练次数的增加,运动组大鼠的游泳速度明显高于对照组（P〈0．01）;另外,运动轨迹的弯曲度表明运动组大鼠还表现出了较强的寻找平台的动机以及对平台位置较为准确的空间定位能力。在空间探索实验中,两组大鼠的游泳速度并没有明显差异,从大鼠在各象限内穿越平台相应位置的次数来看,运动组大鼠在D象限穿越的次数高于对照组,但无统计学差异（P〉0．05）。上述结果提示,低强度的跑台运动在短时间内便可以明显提高幼龄大鼠的学习能力。     

Does metatarsal/femur ratio predict maximal running speed in cursorial mammals?

We tested the hypothesis that hind limb proportions may be used to predict locomotor performance in a sample of 49 species of primarily cursorial mamals. Data on maximal sprint running speeds taken from published sources were related to measurements of hind limb lengths. To control for statistical complications due to the hierarchical nature of phylogenetic relationships, we used Felsenstein's (1985) independent contrasts method for analysing comparative data, and a composite phylogeny for all 49 species, based on a variety of published sources. The independent contrasts method indicates that maximal running speed does not covary significantly with body mass for this sample of mammals (mass range= 2.5–2,000 kg). Even though quality of the available speed data is highly variable, both metatarsal/femur ratio—the traditional index of 'cursoriality' in mammals—and hind limb length (corrected for body size) are significant predictors of maximal running speed. When only fully curorial species are included in the analyses (n = 32), hind limb length still significantly predicts speed (r²= 16%), but MT/F ratio does not. Although ungulates tend to have larger MT/F ratios than do Carnivora, they are not generally faster; relatonships between speed and limb proportions within the two clades show no significant differences. These and previous results suggest that hind limb proportions and maximal running speed may not have evolved in a tightly coupled fashion. Prediction of locomotor performance of extinct forms, based solely on their limb proportions, should be undertaken with caution.     

A movement criterion for running

The adjustment of the leg during running was addressed using a spring-mass model with a fixed landing angle of attack. The objective was to obtain periodic movement patterns. Spring-like running was monitored by a one-dimensional stride-to-stride mapping of the apex height to identify mechanically stable fixed points. We found that for certain angles of attack, the system becomes self-stabilized if the leg stiffness was properly adjusted and a minimum running speed was exceeded. At a given speed, running techniques fulfilling a stable movement pattern are characterized by an almost constant maximum leg force. With increasing speed, the leg adjustment becomes less critical. The techniques predicted for stable running are in agreement with experimental studies. Mechanically self-stabilized running requires a spring-like leg operation, a minimum running speed and a proper adjustment of leg stiffness and angle of attack. These conditions can be considered as a movement criterion for running.     

A little bit faster: Lower extremity joint kinematics and kinetics as recreational runners achieve faster speeds

There appears a linear relationship between small increases in running speed and cardiovascular health benefits. Encouraging or coaching recreational runners to increase their running speed to derive these health benefits might be more effective if their joint level kinematic and kinetic strategy was understood. The aim of this investigation was to compare the peak sagittal plane motions, moments, and powers of the hip, knee and ankle at 85%, 100%, 115% and 130% of self-selected running speed. Overground running data were collected in 12 recreational runners (6 women, 6 men) with a full body marker set using a 12-camera Vicon MX system with an AMTI force plate. Kinematics and kinetics were analyzed with Vicon Nexus software. Participants chose to run at 2.6 ± 0.5 m/s (85%); 3.0 ± 0.5 m/s (100%); 3.3 ± 0.5 m/s (115%); and 3.7 ± 0.5 m/s (130%); these four speeds approximately correspond to 6:24-, 5:33-, 5:03-, and 4:30-min kilometer running paces. Running speed had a significant effect (P < 0.05) on peak kinematic and kinetic variables of the hips, knees and ankles, with peak sagittal hip moments invariant (P > 0.54) and the peak sagittal ankle power generation (P < 0.0001) the most highly responsive variable. The timing of the peak sagittal extensor moments and powers at the hip, knee and ankle were distributed across stance in a sequential manner. This study shows that running speed affects lower limb joint kinematics and kinetics and suggests that specific intersegmental kinetic strategies might exist across the narrow range of running speeds.     

Effects of treadmill activity on cerebellar cyclic GMP

Ovariectomized, female Sprague-Dawley rats were trained to run on a treadmill for up to several minutes. On the day of the experiment the treadmill speed was adjusted to either 680, 1360 or 2720 cm/minute and separate groups of 5 rats were required to run at each of these speeds for either 15, 30, 45 or 60 seconds. Immediately after running on the treadmill, each rat was killed by microwave irradiation, and the cerebellum was collected for subsequent determination of cyclic guanosine monophosphate (cGMP). The time to the onset of an elevation of cGMP and the maximal elevation obtained were functions of the speed of running. The first significant increase in cerebellar cGMP over that observed in inactive control rats occurred within 15 seconds after the onset of running. Regardless of the speed of running, the cGMP content reached a plateau after 45 seconds.     

Groucho running

An important determinant of the mechanics of running is the effective vertical stiffness of the body. This stiffness increases with running speed. At any one speed, the stiffness may be reduced in a controlled fashion by running with the knees bent more than usual. In a series of experiments, subjects ran in both normal and flexed postures on a treadmill. In other experiments, they ran down a runway and over a force platform. Results show that running with the knees bent reduces the effective vertical stiffness and diminishes the transmission of mechanical shock from the foot to the skull but requires an increase of as much as 50% in the rate of O2 consumption. A new dimensionless parameter (u omega 0/g) is introduced to distinguish between hard and soft running modes. Here, omega 0 is the natural frequency of a mass-spring system representing the body, g is gravity, and u is the vertical landing velocity. In normal running, this parameter is near unity, but in deep-flexed running, where the aerial phase of the stride cycle almost disappears, u omega 0/g approaches zero.     

The role of the stretch reflex in the gastrocnemius muscle during human locomotion at various speeds.

In the present study, the fascicle length (L(fa)) of the human medial gastrocnemius (MG) muscle was monitored to evaluate possible input from the short-latency stretch reflex (SLR) during the stance phase of running and to examine its timing at various running speeds. Eight subjects ran at 2.0, 3.5, 5.0, and 6.5 m/s. The L(fa) was measured with the high-speed ultrasound fascicle scanning together with kinematics and myoelectrical activities. The amplitudes and onset latency of SLR activities were determined. During ground contact, the sudden MG fascicle stretch occurred during the early contact at all running speeds. This was followed by the fascicle shortening. The timing of fascicle stretch depended on running speed and type of foot contact. In slower speed conditions (2.0, 3.5, 5 m/s), the MG fascicle stretch and the corresponding SLR activities occurred during the middle of the braking phase. In fast-speed running (6.5 m/s), however, the MG fascicle stretch occurred later compared with the lower speed. The corresponding SLR activities occurred significantly later at the end of the braking phase. In addition to the clear demonstration of the different timings of SLR in MG during ground contact of running, the results imply that the role of the MG SLR during the stance phase of running can be different between fast- and slow-speed running conditions.     

Variations in running activity and enzymatic adaptations in voluntary running rats

The running behavior and biochemical markers of oxidative and glycolytic activities associated with voluntary running activity were studied in male Sprague-Dawley rats after 6 wk of training in exercise wheel cages. Twenty-four-hour recordings of running activity were used to quantify the number of individual running bouts, their duration and running speed, and the distance run per day. We then established three categories of voluntary running activity based on the mean distance run per day during the last 3 wk of training: low-activity runners averaged 2-5 km/day, medium runners 6-9 km/day, and high runners greater than 11 km/day. Each group demonstrated an intermittent, nocturnal running pattern, at relatively high intensities, with a similar mean running speed for all groups (avg approximately 45 m/min). Differences in total distance run per day were the result of variations in both the number and duration of individual running bouts. Specifically, high runners (n = 7) had 206 +/- 30 individual running bouts per 24 h, each lasting 87 +/- 7 s; medium runners (n = 7) 221 +/- 22 running bouts, lasting 47 +/- 5 s; and low runners (n = 7) 113 +/- 7 bouts, each lasting 40 +/- 7 s. Voluntary running depressed the rate of body weight gain compared with sedentary control rats, despite an increased food and water intake for all runners. Furthermore, drinking activity was temporally associated with running periods.(ABSTRACT TRUNCATED AT 250 WORDS)     

Ontogenetic and individual variation in size, shape and speed in the Australian agamid lizard Amphibolurus nuchalis

The present study investigates relationships among size, shape and speed in the Australian agamid lizard Amphibolurus nuchalis . Maximal running speed, body mass, snout-vent length, tail length, fore- and hind limb spans and thigh muscle mass were measured in 68 field-fresh individuals spanning the entire ontogenetic size range (1.3 48 g). Relative lengths of both foreand hind limbs decrease with increasing body mass (= negative allometry), whereas relative tail length and thigh muscle mass increase with body mass (= positive allometry). Repeatable and significant differences in maximal running speed exist among individuals. Maximal running speed scales as (body mass)^0.161, and 59% of the variation in maximal speed was related to body mass. Based on the results of the present and previous studies, data on scaling of body proportions alone appear inadequate to infer scaling relationships of functional characters such as top speed.
Surprisingly, individual variation in maximal speed is not related to individual variation in shape (relative limb, tail and body lengths). These components of overall shape are not independent; individuals tended to have either relatively long or relatively short limbs, tails and bodies for their body mass. Even the significant difference in multivariate shape between adult males and females has no measurable consequences for maximal speed. Speeds of field-fresh animals did not vary on a seasonal basis, and eight weeks of captivity had no effect on maximal running speeds. Gravid females and long-term (obese) captive lizards were both approximately 12% slower than field-fresh lizards.     

Rat muscle blood flows during high-speed locomotion

We previously studied blood flow distribution within and among rat muscles as a function of speed from walking (15 m/min) through galloping (75 m/min) on a motor-driven treadmill. The results showed that muscle blood flows continued to increase as a function of speed through 75 m/min. The purpose of the present study was to have rats run up to maximal treadmill speeds to determine if blood flows in the muscles reach a plateau as a function of running speed over the animals' normal range of locomotory speeds. Muscle blood flows were measured with radiolabeled microspheres at 1 min of running at 75, 90, and 105 m/min in male Sprague-Dawley rats. The data indicate that even at these relatively high treadmill speeds there was still no clear evidence of a plateau in blood flow in most of the hindlimb muscles. Flows in most muscles continued to increase as a function of speed. These observed patterns of blood flow vs. running speed may have resulted from the rigorous selection of rats that were capable of performing the high-intensity exercise and thus only be representative of a highly specific population of animals. On the other hand, the data could be interpreted to indicate that the cardiovascular potential during exercise is considerably higher in laboratory rats than has normally been assumed and that inadequate blood flow delivery to the muscles does not serve as a major limitation to their locomotory performance.     

Repeated sprint training improves intermittent peak running speed in team-sport athletes

The aim of this study was to compare the effect of 2 repeated sprint training interventions on an intermittent peak running speed (IPRS) test designed for Australian Rules football. The test required participants to perform 10 × 10-m maximal efforts on an 80-m course every 25 seconds, for each of which the mean peak speed (kilometers per hour) was recorded to determine IPRS. The training interventions were performed twice weekly for 4 weeks immediately before regular football training. In the constant volume intervention (CVol), sprint repetition number remained at 10 (n = 9), and in the linear increase in volume (LIVol) intervention, repetition number increased linearly each week by 2 repetitions (n = 12). Intermittent peak running speed, 300-m shuttle test performance, and peak running speed were assessed before and upon completion of training. All measures were compared to a control group (CON; n = 8) in which players completed regular football training exclusively. Intermittent peak running speed performance in CVol and LIVol improved significantly (p < 0.01) by 5.2 and 3.8%, respectively, with no change in IPRS for CON. There were no differences in IPRS changes between CVol and LIVol. Additionally, peak running speed improved significantly (p < 0.01) by 5.1% for CVol, whereas 300-m shuttle performance improved significantly (p < 0.01) by 2.6% for LIVol only. Intermittent peak running speed, 300-m shuttle performance and peak running speed were improved after 4 weeks of training; however, progressively increasing sprint repetition number had no greater advantage on IPRS adaptation. Additionally, exclusive regular football training over a 4-week period is unlikely to improve IPRS, peak running speed, or 300-m shuttle performance.     

Stretching Your Energetic Budget: How Tendon Compliance Affects the Metabolic Cost of Running

Muscles attach to bones via tendons that stretch and recoil, affecting muscle force generation and metabolic energy consumption. In this study, we investigated the effect of tendon compliance on the metabolic cost of running using a full-body musculoskeletal model with a detailed model of muscle energetics. We performed muscle-driven simulations of running at 2–5 m/s with tendon force–strain curves that produced between 1 and 10% strain when the muscles were developing maximum isometric force. We computed the average metabolic power consumed by each muscle when running at each speed and with each tendon compliance. Average whole-body metabolic power consumption increased as running speed increased, regardless of tendon compliance, and was lowest at each speed when tendon strain reached 2–3% as muscles were developing maximum isometric force. When running at 2 m/s, the soleus muscle consumed less metabolic power at high tendon compliance because the strain of the tendon allowed the muscle fibers to operate nearly isometrically during stance. In contrast, the medial and lateral gastrocnemii consumed less metabolic power at low tendon compliance because less compliant tendons allowed the muscle fibers to operate closer to their optimal lengths during stance. The software and simulations used in this study are freely available at simtk.org and enable examination of muscle energetics with unprecedented detail.     

The consistency of maximum running speed measurements in humans using a feedback-controlled treadmill,and a comparison with maximum attainable speed during overground locomotion

Consistent measurement of maximum running speed overground is problematic due to the difficulty in precise, continual measurement of speed, and the substantial workload in accelerating the body promoting the onset of fatigue. Treadmills remove the requirement for acceleration which enables more repeats. They also allow experiments to be carried out in controlled environments and where space is limited, but they usually depend on manual and subjective speed control. Here we used a draw-wire position sensor and a proportional–derivative (PD) controller to automatically adjust treadmill belt speed of a large equine treadmill. The feedback loop took the real-time position and velocity of the runner relative to the front of the treadmill as input. This control system allowed runners to accelerate from walking speed to a peak running speed within a few strides and then decelerate as quickly as they wished. We used the system to evaluate the variation in maximum speed determination that results from one trial to 10 trials, in eleven individuals. Three trials gave a maximum speed 97.8% of that achieved after ten. The approach used is appropriate for any treadmill where the running zone length is greater than three metres and the speed controller can be externally controlled. Subjects ran 11.5% faster on the treadmill than overground, part of which can be explained by the removal of aerodynamic drag and the fatigue of overground running. Additional factors may, however, contribute to athletes running faster on a treadmill, for instance some aspect of stability or control.     

Grip and limb force limits to turning performance in competition horses

Manoeuverability is a key requirement for successful terrestrial locomotion, especially on variable terrain, and is a deciding factor in predator-prey interaction. Compared with straight-line running, bend running requires additional leg force to generate centripetal acceleration. In humans, this results in a reduction in maximum speed during bend running and a published model assuming maximum limb force as a constraint accurately predicts how much a sprinter must slow down on a bend given his maximum straight-line speed. In contrast, greyhounds do not slow down or change stride parameters during bend running, which suggests that their limbs can apply the additional force for this manoeuvre. We collected horizontal speed and angular velocity of heading of horses while they turned in different scenarios during competitive polo and horse racing. The data were used to evaluate the limits of turning performance. During high-speed turns of large radius horizontal speed was lower on the bend, as would be predicted from a model assuming a limb force limit to running speed. During small radius turns the angular velocity of heading decreased with increasing speed in a manner consistent with the coefficient of friction of the hoof-surface interaction setting the limit to centripetal force to avoid slipping.     

Assessment of human locomotion by using an insole measurement system and artificial neural networks

A new method for measuring and characterizing free-living human locomotion is presented. A portable device was developed to objectively record and measure foot-ground contact information in every step for up to 24h. An artificial neural network (ANN) was developed to identify the type and intensity of locomotion. Forty subjects participated in the study. The subjects performed level walking, running, ascending and descending stairs at slow, normal and fast speeds determined by each subject, respectively. The device correctly identified walking, running, ascending and descending stairs (accuracy 98.78%, 98.33%, 97.33%, and 97.29% respectively) among different types of activities. It was also able to determine the speed of walking and running. The correlation between actual speed and estimated speed is 0.98, p< 0.0001. The average error of walking and running speed estimation is -0.050+/-0.747 km/h (mean +/- standard deviation). The study has shown the measurement of duration, frequency, type, and intensity of locomotion highly accurate using the new device and an ANN. It provides an alternative tool to the use of a gait lab to quantitatively study locomotion with high accuracy via a small, light and portable device, and to do so under free-living conditions for the clinical applications.     

Comparison of energy expenditure and substrate oxidation between walking and running in men and women

[Purpose]The present study compared energy metabolism between walking and running at equivalent speeds during two incremental exercise tests.[Methods]Thirty four university students (18 males, 16 females) were recruited. Each participant completed two trials, consisting of walking (Walk) and running (Run) trials on different days, with 2-3 days apart. Exercise on a treadmill was started from initial stage of 3 min (3.0 k/m in Walk trial, 5.0 km/h in Run trial), and the speed for walking and running was progressively every minute by 0.5 km/h. The changes in metabolic variables, heart rate (HR), and rating of perceived exertion (RPE) during exercise were compared between the trials.[Results]Energy expenditure (EE) increased with speed in each trial. However, the Walk trial had a significantly higher EE than the Run trial at speeds exceeding 92 ± 2 % of the maximal walking speed (MWS, p < 0.01). Similarly, carbohydrate (CHO) oxidation was significantly higher in the Walk trial than in the Run trial at above 92 ± 2 %MWS in males (p < 0.001) and above 93 ± 1 %MWS in females (p < 0.05).[Conclusion]These findings suggest that EE and CHO oxidation during walking increase non-linearly with speed, and walking at a fast speed causes greater metabolic responses than running at the equivalent speed in young participants.     

Effects of size, sex, and voluntary running speeds on costs of locomotion in lines of laboratory mice selectively bred for high wheel-running activity

Selective breeding for over 35 generations has led to four replicate (S) lines of laboratory house mice (Mus domesticus) that run voluntarily on wheels about 170% more than four random-bred control (C) lines. We tested whether S lines have evolved higher running performance by increasing running economy (i.e., decreasing energy spent per unit of distance) as a correlated response to selection, using a recently developed method that allows for nearly continuous measurements of oxygen consumption (VO2) and running speed in freely behaving animals. We estimated slope (incremental cost of transport [COT]) and intercept for regressions of power (the dependent variable, VO2/min) on speed for 49 males and 47 females, as well as their maximum VO2 and speeds during wheel running, under conditions mimicking those that these lines face during the selection protocol. For comparison, we also measured COT and maximum aerobic capacity (VO2max) during forced exercise on a motorized treadmill. As in previous studies, the increased wheel running of S lines was mainly attributable to increased average speed, with males also showing a tendency for increased time spent running. On a whole-animal basis, combined analysis of males and females indicated that COT during voluntary wheel running was significantly lower in the S lines (one-tailed P=0.015). However, mice from S lines are significantly smaller and attain higher maximum speeds on the wheels; with either body mass or maximum speed (or both) entered as a covariate, the statistical significance of the difference in COT is lost (one-tailed P> or =0.2). Thus, both body size and behavior are key components of the reduction in COT. Several statistically significant sex differences were observed, including lower COT and higher resting metabolic rate in females. In addition, maximum voluntary running speeds were negatively correlated with COT in females but not in males. Moreover, males (but not females) from the S lines exhibited significantly higher treadmill VO2max as compared to those from C lines. The sex-specific responses to selection may in part be consequences of sex differences in body mass and running style. Our results highlight how differences in size and running speed can account for lower COT in S lines and suggest that lower COT may have coadapted in response to selection for higher running distances in these lines.     

Maximal oxygen uptake during treadmill walking and running at various speeds.

Three groups of male subjects, average fitness (AF, N = 12), high fitness (HF, N = 7) and highly fit competitive race walkers (CRW, N = 3) performed maximal treadmill tests walking at 3.5 and 4.5 mph and running at 4.5, 5.5, 7.0, and 8.5 mph. In addition, the HF group performed a running test at 10.0 mph and the CRW group performed a walking test at 5.5 mph. All maximal oxygen uptake (VO2 max) tests with the exception of the 3.5 mph walking test (modified Balke test) were discontinuous in nature. VO2 max obtained from walking tests was similar regardless of speed within each group. Walking VO2 max was significantly lower than running VO2 max which was found to be similar over a speed range of 4.5 to 8.5 mph in the AF group. Running at 4.5 mph (HF group) and 4.5 and 5.5 mph (CRW group) resulted in lower VO2 max levels than running at speeds greater than or equal to 7.0 mph. Associated physiological variables (heart rate, ventilation, and respiratory exchange ratio) did not demonstrate a discernable pattern with reference to mode of locomotion (walking versus running) or speed. It was concluded that VO2 max elicited during walking is independent of speed and less than VO2 max obtained during running. Running VO2 max was interrelated with speed of running and state of training.     

Locomotion in terrestrial mammals: the influence of body mass, limb length and bone proportions on speed

Traditionally a few limb proportions or total limb lengths have been regarded as indicative of peak running velocity. This is due to physical principles (inferred in- and outvelocities around the joints, stride lengths) and also the observation that fast-moving animals tend to share a number of purported key features which are either absent or not developed to near the same extent in slower moving forms. Previous studies have shown hind limb length and metatarsus/femur ratio to be correlated significantly, albeit modestly with running speed. These studies have nearly all been bivariate analyses. Based on the physical principles, there is reason to suppose that more variables than just m/f ratio could be important as adaptations for fast locomotion, and also that bivariate analyses are too simple. In this study a sample of 76 running mammals was used, with running speeds taken from literature. A number of osteological parameters were discovered to covary significantly with peak running speed, albeit only modestly. Using the information from phylogeny reduced all correlations, often significantly so. Multivariate analyses resulted in markedly higher correlation coefficients. Animals probably do not optimize their anatomy for the purpose of running very fast, which occurs only on rare occasions, but for reducing costs of locomotion.  © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society , 2002, 136 , 685–714.