Oxygen delivery does not limit peak running speed during incremental downhill running to exhaustion

Oxygen consumption (VO2), ventilation (VI), respiratory exchange ratio (R), stride frequency and blood lactate concentrations were measured continuously in nine trained athletes during two continuous incremental treadmill runs to exhaustion on gradients of either 0 degree or -3 degrees. Compared to the run at 0 degree gradient, the athletes reached significantly higher maximal treadmill velocities but significantly lower VO2, VI, R and peak blood lactate concentrations (P less than 0.001) during downhill running. These lower VO2 and blood lactate concentrations at exhaustion indicated that factors other than oxygen delivery limited maximal performance during the downhill run. In contrast, stride frequencies were similar at each treadmill velocity; the higher maximal speed during the downhill run was achieved with a significantly longer stride length (P less than 0.001); maximal stride frequency was the same between tests. Equivalent maximal stride frequencies suggested that factors determining the rate of lower limb stride recovery may have limited maximal running speed during downhill running and, possibly, also during horizontal running.     

Explaining the scaling of transport costs: the role of stride frequency and stride length

The mechanisms which enable large animals to transport a unit of body mass through a unit distance at a lower metabolic cost than smaller animals have been the subject of numerous studies. Recent investigations have concluded that stride frequency is a main determinant. We examine the role of both stride frequency and stride length in determining the scaling of the cost of transport.
Slopes for regressions between stride frequency and speed and stride length and speed were determined in four species of rodents. These data were pooled with literature values for the slopes of stride frequency, stride length and cost of locomotion (all vs. speed) for a total of 17 species ranging in size from 30 g to 250 kg. Interspecific equations were calculated for each of these slopes versus body mass, and residuals from these allometric lines were calculated. Residuals were compared to see if variation in the rate of cost increase at a given size is related to variation in the rates of stride frequency and/or stride length increase.
The residual analysis revealed that the variation in transport cost is explicable only in terms of the interaction of stride frequency and stride length slopes. The product of the scaling exponents for stride frequency slope and stride length slope is not significantly different from the scaling exponent for the cost of transport. A model seeking to explain the scaling of the cost of transport must therefore consider the influence of both stride length and stride frequency.
We propose that absolutely longer limbs allow large animals to minimize the rate of increase of stride frequency and stride length with speed, and that this allows utilization of muscles with lower intrinsic rates of contraction, which in turn results in a lower mass-specific cost of transport.     

Steps to Take to Enhance Gait Stability: The Effect of Stride Frequency,Stride Length,and Walking Speed on Local Dynamic Stability and Margins of Stability

The purpose of the current study was to investigate whether adaptations of stride length, stride frequency, and walking speed, independently influence local dynamic stability and the size of the medio-lateral and backward margins of stability during walking. Nine healthy subjects walked 25 trials on a treadmill at different combinations of stride frequency, stride length, and consequently at different walking speeds. Visual feedback about the required and the actual combination of stride frequency and stride length was given during the trials. Generalized Estimating Equations were used to investigate the independent contribution of stride length, stride frequency, and walking speed on the measures of gait stability. Increasing stride frequency was found to enhance medio-lateral margins of stability. Backward margins of stability became larger as stride length decreased or walking speed increased. For local dynamic stability no significant effects of stride frequency, stride length or walking speed were found. We conclude that adaptations in stride frequency, stride length and/or walking speed can result in an increase of the medio-lateral and backward margins of stability, while these adaptations do not seem to affect local dynamic stability. Gait training focusing on the observed stepping strategies to enhance margins of stability might be a useful contribution to programs aimed at fall prevention.     

The entrainment of breathing rhythm to stride frequency in the dik-dik antelope (Rhynchotragus kirki) with observations on the thermoregulatory consequences

1. Three adult dik-dik antelopes with an average weight of 4.5 kg were used to investigate the rate of oxygen consumption (VO2) during exercise and the entrainment of respiratory rate with stride frequencies at running velocities between 2-11 km/hr. 2. The results of VO2 and the metabolic cost of horizontal locomotion were found to agree with what would be expected of an animal of this size. 3. Respiratory frequencies recorded during the most strenuous exercise were approximately 50% below the value observed when dik-diks are exposed to an ambient temperature (Ta) of 42 degrees C. 4. Respiratory evaporative heat loss was estimated to account for only 4% of the total heat production during exercise. 5. Respiratory frequencies were found to be entrained quite strongly to stride frequencies. The thermoregulatory consequences of this entrainment is discussed.     

Effect of locomotor respiratory coupling on respiratory evaporative heat loss in the sheep.

During galloping, many animals display 1:1 coupling of breaths and strides. Locomotor respiratory coupling (LRC) may limit respiratory evaporative heat loss (REHL) by constraining respiratory frequency (f). Five sheep were exercised twice each, according to a five-step protocol: 5 min at the walk, 5 min at the trot (trot1), 10 min at the gallop, 5 min at the trot (trot2), and 5 min at the walk. Rectal temperature (T(re)), stride frequency, f, REHL, and arterial CO(2) tension and pH were measured at each step. Tidal volume (VT) was calculated. LRC was observed only during galloping. The coupling ratio remained at 1:1 while VT increased continuously during galloping, causing REHL to increase from 2.9 +/- 0.2 (SE) W/kg at the end of trot1 to a peak of 5.3 +/- 0.3 W/kg. T(re) rose from 39.0 +/- 0.1 degrees C preexercise to 40.2 +/- 0.2 degrees C at the end of galloping. At the gallop-trot2 transition, VT fell and f rose, despite a continued rise in T(re). Arterial CO(2) tension fell from 36.5 +/- 1.1 Torr preexercise to 31.8 +/- 1.4 Torr by the end of trot1 and then further to 21.5 +/- 1.2 Torr by the end of galloping, resulting in alkalosis. In conclusion, LRC did not prevent increases in REHL in sheep because VT increased. The increased VT caused hypocapnia and presumably elevated the cost of breathing.     

The effect of uphill stride manipulation on race walking gait

Stride length analysis represents an easy method for assessing race walking kinematics. However, the stride parameters emerging from such an analysis have never been used to design a training protocol aimed at increasing stride length. With this aim, we investigated the effects of stride frequency manipulation during three weeks of uphill (2%) training on stride length at iso-efficiency speed. Twelve male race walkers were randomly allocated to one of two training groups: stride frequency manipulation (RWM, n=6) and free stride frequency (RWF, n=6). Results. Kinematic parameters measured before and after the 3-week training in RWM showed increased stride length (4.54%; p<0.0001) and contact time (4.58%; p<0.001); inversely, a decreased stride frequency (4.44%; p<0.0001) and internal work (7.09%; p<0.05) were found. In RWF the effect of the training showed a decrease in stride length (1.18%; p<0.0001) and contact time (<1%; p<0.0001) with respect to baseline conditions and an increased stride frequency and internal work of 1.19% (p<0.0001). These results suggest that using slopes (2%) as RWM could help coaches to provide some training methods that would improve an athlete''s performance, through increasing stride length without altering his or her race walking technique or metabolic demands.     

The effect of stride length on the dynamics of barefoot and shod running

A number of interventions and technique changes have been proposed to attempt to improve performance and reduce the number of running related injuries. Running shoes, barefoot running and alterations in spatio-temporal parameters (stride frequency and stride length) have been associated with significant kinematic and kinetic changes, which may have implications for performance and injury prevention. However, because footwear interventions have been shown to also affect spatio-temporal parameters, there is uncertainty regarding the origin of the kinematic and kinetic alterations. Therefore, the purpose of this study was to independently evaluate the effects of shoes and changes in stride length on lower extremity kinetics. Eleven individuals ran over-ground at stride lengths ±5 and 10% of their preferred stride length, in both the barefoot and shod condition. Three-dimensional motion capture and force plate data were captured synchronously and used to compute lower extremity joint moments. We found a significant main effect of stride length on anterior–posterior and vertical GRFs, and sagittal plane knee and ankle moments in both barefoot and shod running. When subjects ran at identical stride lengths in the barefoot and shod conditions we did not observe differences for any of the kinetic variables that were measured. These findings suggest that barefoot running triggers a decrease in stride length, which could lead to a decrease in GRFs and sagittal plane joint moments. When evaluating barefoot running as a potential option to reduce injury, it is important to consider the associated change in stride length.     

Hemiplegic gait: a kinematic analysis using walking speed as a basis.

The kinematics of treadmill ambulation of stroke patients (N = 9) and healthy subjects (N = 4) was studied at a wide range of different velocities (i.e. 0.25-1.5 m s-1), with a focus on the transverse rotations of the trunk. Video recordings revealed, for both stroke patients and healthy subjects, similar relations between walking speed and stride length as well as stride frequency. The phase difference between pelvic and thoracic rotations (i.e. trunk rotation) and the total range of trunk rotation were almost linearly related to the walking speed. Healthy subjects showed a marked increase in pelvic rotation from 1 to 1.5 m s-1. Using dimensional analysis in a comparison between stroke patients and healthy subjects, invariances in the coordination of gait were found for stride length, stride frequency, pelvic rotation, and trunk rotation. Constant relations were obtained between, on the one hand, dimensionless velocity and, on the other, dimensionless stride length as well as stride frequency. Transitions were found between the velocities 0.75 and 1 m s-1 for dimensionless pelvic rotation and trunk rotation, indicating that, from this velocity range onwards, pelvic swing lengthens the stride: rotations of pelvis, thorax and trunk become tightly coordinated. On the basis of the dimensionless stride length, stride frequency, pelvic rotation and trunk rotation, deficits in the gait of stroke patients could be quantified. It is concluded that walking speed is an important control parameter, which should be used as a basic variable in the evaluation of the gait of stroke patients.     

How do load carriage and walking speed influence trunk coordination and stride parameters?

To determine the effects of load carriage and walking speed on stride parameters and the coordination of trunk movements, 12 subjects walked on a treadmill at a range of walking speeds (0.6-1.6 m s(-1)) with and without a backpack containing 40% of their body mass. It was hypothesized that compared to unloaded walking, load carriage decreases transverse pelvic and thoracic rotation, the mean relative phase between pelvic and thoracic rotations, and increases hip excursion. In addition, it was hypothesized that these changes would coincide with a decreased stride length and increased stride frequency. The findings supported the hypotheses. Dimensionless analyses indicated that there was a significantly larger contribution of hip excursion and smaller contribution of transverse plane pelvic rotation to increases in stride length during load carriage. In addition, there was a significant effect of load carriage on the amplitudes of transverse pelvic and thoracic rotation and the relative phase of pelvic and thoracic rotation. It was concluded that the shorter stride length and higher stride frequency observed when carrying a backpack is the result of decreased pelvic rotation. During unloaded walking, increases in pelvic rotation contribute to increases in stride length with increasing walking speed. The decreased pelvic rotation during load carriage requires an increased hip excursion to compensate. However, the increase in hip excursion is insufficient to fully compensate for the observed decrease in pelvis rotation, requiring an increase in stride frequency during load carriage to maintain a constant walking speed.     

Effect of stride frequency on thermoregulatory responses during endurance running in distance runners

Changing stride frequency may influence oxygen uptake and heart rate during running as a function of running economy and central command. This study investigated the influence of stride frequency manipulation on thermoregulatory responses during endurance running. Seven healthy endurance runners ran on a treadmill at a velocity of 15 km/h for 60 min in a controlled environmental chamber (ambient temperature 27 °C and relative humidity 50%), and stride frequency was manipulated. Stride frequency was intermittently manipulated by increasing and decreasing frequency by 10% from the pre-determined preferred frequency. These periods of increase or decrease were separated by free frequency running in the order of free stride frequency, stride frequency manipulation (increase or decrease), free stride frequency, and stride frequency manipulation (increase or decrease) for 15 min each. The increased and decreased stride frequencies were 110% and 91% of the free running frequency, respectively (196±6, 162±5, and 178±5 steps/min, respectively, P<0.01). Compared to the control, stride frequency manipulation did not affect rectal temperature, heart rate, or the rate of perceived exhaustion during running. Whole-body sweat loss increased significantly when stride frequency was manipulated (1.48±0.11 and 1.57±0.11 kg for control and manipulated stride frequencies, respectively, P<0.05), but stride frequency had a small effect on sweat loss overall (Cohen's d=0.31). A higher mean skin temperature was also observed under mixed frequency conditions compared to that in the control (P<0.05). While the precise mechanisms underlying these changes remain unknown (e.g. running economy or central command), our results suggest that manipulation of stride frequency does not have a large effect on sweat loss or other physiological variables, but does increase mean skin temperature during endurance running.     

Hindlimb morphology and locomotor performance in waders: an evolutionary approach

Locomotion performance (measured as stride frequency and stride length) was studied in 16 species of waders. Differences in hindlimb morphology (osteology and myology) were analysed among species. Evolutionary changes in both locomotion and morphological variables were analysed using comparative methods revealing the existence of some ecomorphological patterns relating these two sets of characters. Evolutionary changes in stride frequency were correlated with changes in the muscles M. iliotibialis cranialis, M. iliotibiales lateralis and M. gastrocnemius, whereas changes in stride length showed correlated evolution with changes in the length of distal segments of the leg. We identify two different evolutionary strategies in locomotion of waders. One is a change in distal leg segments (skeletal system), an adaptive modification that increases stride length; the second is a change in the skeletal-muscular system, providing an increase in muscular performance (force or speed of contraction) in several muscles, and is an adaptation that increases stride frequency.     

Lower-extremity ground reaction forces in collegiate baseball pitchers

The purpose of this study was to investigate ground reaction forces (GRF) in collegiate baseball pitchers and their relationship to pitching mechanics. Fourteen healthy collegiate baseball pitchers participated in this study. High-speed video and force plate data were collected for fastballs from each pitcher. The average ball speed was 35 ± 3 m/sec (78 ± 7 mph). Peak GRFs of 245 ± 20% body weight (BW) were generated in an anterior or braking direction to control descent. Horizontal GRFs tended to occur in a laterally directed fashion, reaching a peak of 45 ± 63% BW. The maximum vertical GRF averaged 202 ± 43% BW approximately 45 milliseconds after stride foot contact. A correlation between braking force and ball velocity was evident. Because of the downward inclination and rotation of the pitching motion, in addition to volume, shear forces may occur in the musculoskeletal tissues of the stride limb leading to many of the lower-extremity injuries seen in this athletic population.     

Axial-Appendicular Dynamics and the Integration of Breathing and Gait in Mammal

SYNOPSIS. This paper proposes a biomechanical model for locomotor-respiratorycoupling (LRC) in galloping mammals in which gait and breathingcycles are phase-locked on a 1:1 basis. It also explores someof the physiological and neuromotor implications of LRC. The mechanical coupling of locomotor and respiratory cyclesdepends upon the coordinated, reciprocal oscillations of thecranio-cervical and lumbo-pelvic components of the axial systemand their attendant actions on the intervening thorax via muscularlinkages. Concurrently, accelerational and decelerational forcesimparted to the axial system by the limbs help to drive lungventilation by inducing inertial displacements of a "visceralpiston’ connected to the diaphragm. Several lines of evidence(including cineradiographic data) suggest that an importantfunction of the crural diaphragm is to control the displacementof the visceral piston. The kinematics of LRC indicate thatthe interosseous intercostal muscles must simultaneously operateto assure thoracic stability against locomotor stresses as wellas to promote breathing. The former may be their more essentialrole, however. The characteristic design of the rib cage incursorial mammals (=deep and narrow) appears to maximize theleverage of certain "accessory respiratory muscles" (i.e., sternocleidomastoid,scalenes) while minimizing torsional loading of the thorax duringforelimb support. Physiological implications of LRC include the prediction thatlarge mammals will breathe relatively faster and with relativelysmaller lung volumes when galloping than small species. An additionalprediction, that running mammals could automatically gear lungventilation to speed by simply linking breathing rate to stridefrequency and depth of breath (=tidal volume) to stride length,appears to be supported by experimental data from horses. Finally,the neuromotor basis of LRC probably depends upon the directinteraction of central pattern generators for locomotion andrespiration. This interaction might be modulated, however, byafferent input from thoracic mechanoreceptors, particularlythe intercostal stretch receptors.     

Stride length and its determinants in humans, early hominids, primates, and mammals

Primate stride lengths during quadrupedal locomotion are very long when compared to those of nonprimate quadrupedal mammals at the speed of trot/gallop transition. These exceptional lengths are a consequence of the relatively long limbs of primates and the large angular excursions of their limbs during quadrupedalism. When quadrupedal primates employ bipedal gaits they exhibit much lower angular excursions. Consequently their bipedal stride lengths do not appear to be exceptional in length when compared to other mammals. Angular excursions of the lower limbs of modern humans are not exceptionally large. However, when running, humans exhibit relatively long periods of flight (i.e., they have low duty factors) when compared to other mammals including primates. Because of these long periods of flight and their relative long lower limbs, humans have running stride lengths that are at the lower end of the range of stride lengths of quadrupedal primates. The stride length of the Laetoli hominid trails are evaluated in this context.     

The Laetoli footprints and early hominin locomotor kinematics

A critical question in human evolution is whether the earliest bipeds walked with a bent-hip, bent-knee gait or on more extended hindlimbs. The differences between these gaits are not trivial, because the adoption of either has important implications for the evolution of bipedalism. In this study, we re-examined the Laetoli footprints to determine whether they can provide information on the locomotor posture of early hominins. Previous researchers have suggested that the stride lengths of Laetoli hominins fall within the range of modern human stride lengths and therefore, Laetoli hominins walked with modern-human-like kinematics. Using a dynamic-similarity analysis, we compared Laetoli hominin stride lengths with those of both modern humans and chimpanzees. Our results indicate that Laetoli hominins could have used either a bent-hip, bent-knee gait, similar to a chimpanzee, or an extended-hindlimb gait, similar to a human. In fact, our data suggest that the Laetoli hominins could have walked near their preferred speeds using either limb posture. This result contrasts with most previous studies, which suggest relatively slow walking speeds for these early bipeds. Despite the many attempts to discern limb-joint kinematics from Laetoli stride lengths, our study concludes that stride lengths alone do not resolve the debate over early hominin locomotor postures.     

Symmetrical and asymmetrical gaits in the mouse: patterns to increase velocity

The gaits of the adult SWISS mice during treadmill locomotion at velocities ranging from 15 to 85 cm s^–1 have been analysed using a high-speed video camera combined with cinefluoroscopic equipment. The sequences of locomotion were analysed to determine the various space and time parameters of limb kinematics. We found that velocity adjustments are accounted for differently by the stride frequency and the stride length if the animal showed a symmetrical or an asymmetrical gait. In symmetrical gaits, the increase of velocity is provided by an equal increase in the stride length and the stride frequency. In asymmetrical gaits, the increase in velocity is mainly assured by an increase in the stride frequency in velocities ranging from 15 to 29 cm s^–1. Above 68 cm s^–1, velocity increase is achieved by stride length increase. In velocities ranging from 29 to 68 cm s^–1, the contribution of both variables is equal as in symmetrical gaits. Both stance time and swing time shortening contributed to the increase of the stride frequency in both gaits, though with a major contribution from stance time decrease. The pattern of locomotion obtained in a normal mouse should be used as a template for studying locomotor control deficits after lesions or in different mutations affecting the nervous system.     

Locomotion patterns in two South American gymnophthalmid lizards: Vanzosaura rubricauda and Procellosaurinus tetradactylus

We quantified gait and stride characteristics (velocity, frequency, stride length, stance and swing duration, and duty factor) in the bursts of locomotion of two small, intermittently moving, closely related South American gymnophthalmid lizards: Vanzosaura rubricauda and Procellosaurinus tetradactylus. They occur in different environments: V. rubricauda is widely distributed in open areas with various habitats and substrates, while P. tetradactylus is endemic to dunes in the semi-arid Brazilian Caatinga. Both use trot or walking trot characterised by a lateral sequence. For various substrates in a gradient of roughness (perspex, cardboard, sand, gravel), both species have low relative velocities in comparison with those reported for larger continuously moving lizards. To generate velocity, these animals increase stride frequency but decrease relative stride length. For these parameters, P. tetradactylus showed lower values than V. rubricauda. In their relative range of velocities, no significant differences in stride length and frequency were recorded for gravel. However, the slopes of a correlation between velocity and its components were lower in P. tetradactylus on cardboard, whereas on sand this was only observed for velocity and stride length. The data showed that the difference in rhythmic parameters between both species increased with the smoothness of the substrates. Moreover, P. tetradactylus shows a highly specialised locomotor strategy involving lower stride length and frequency for generating lower velocities than in V. rubricauda. This suggests the evolution of a central motor pattern generator to control slower limb movements and to produce fewer and longer pauses in intermittent locomotion.     

Respiratory reactions on microinjections of GABA and baclofen into the Bötzinger complex and the pre-Bötzinger complex in rats

In adult anaesthetized rats the respiratory reactions to microinjections of GABA (10(-5) M) and baclofen (10(-6) M) into Botzinger complex (BC) and pre-Botzinger complex (PBC) were investigated. It was shown, that GABA microinjections into BC shortened inspiratory time and extended expiratory time while respiratory rate was not changed essentially, under this conditions the tidal volume and ventilation were increased. GABA microinjections into PBC significantly inhibited respiratory rhythm due to inspiratory and expiratory time prolongations and reduced tidal volume. The microinjections of baclofen into BC reduced expiration time and ventilation, and increased respiratory frequency whereas microinjections into PBC increased tidal volume without respiratory rate and expiratory time changes. It is suggested that the reactions observed demonstrate the various contribution of GABAergic mechanisms, including GABA(B)-receptors within BC and PBC, in control of respiratory pattern parameters.     

Lung volume changes in response to altered breathing gas pressure during upright immersion

Since elastic and flow-resistive respiratory work are volume dependent, changes in lung volume during immersion affect respiratory effort. This investigation examined changes in lung volume with air delivery pressure modifications during upright immersion. Static pressure-volume relaxation relationships and lung volumes were obtained from ten immersed subjects breathing air at four delivery pressures: mouth pressure, lung centroid pressure (PLC), and 0.98 kPa above and below PLC. The PLC is the static lung pressure which returns the respiratory relaxation volume (VR) to normal and was previously determined to be +1.33 kPa relative to pressure at the sternal notch. Lung volume changes observed when breathing air at mouth pressure were reversed when air was supplied at PLC. The expiratory reserve volume (ERV) and VR were reduced by 58% and 87%, respectively, during uncompensated immersion. These differences indicated an active defence of ERV and implied that additional static respiratory work was required to overcome transrespiratory pressure gradients.