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.     

An application of cineradiography to the study of locomotion in primates

The cineradiographic technique, whereby, at 64 frames/second, the successive positions of every bone segment of animals in motion can be accurately analyzed, was used to study the vertical leap of a Galago alleni from the ground position. This jump, 13 to 14 times the body length of the animal excluding the tail, is a record among primates challenged only by the tarsier. Such vertical leaps from the ground are a common mode of locomotion for the galago. Two different patterns were observed, a symmetrical and an asymmetrical jump. In the former, the two hind limbs move in the same manner and simultaneously; and the propulsive force is equally distributed in both limbs. In the latter (more frequent in our records), the animal shifts its weight onto one foot during the preparatory phase and lifts the other; thus, only one of the hind limbs is responsible for the launching. The angular variations of every knee and ankle articulation and the successive positions of the hind limb bones were measured and diagrammatically analyzed frame by frame. A short film was made to illustrate the cineradiographic technique; it cinematographically and cineradiographically records in successive sequence the galago's leap from the ground. The symmetrical and asymmetrical jumps are also schematically presented.     

Symmetry perception by poultry chicks and its implications for three-dimensional object recognition

Bilateral symmetry is visually salient to diverse animals including birds, but whereas experimental studies typically use bilaterally symmetrical two-dimensional patterns that are viewed approximately fronto-parallel; in nature, animals observe three-dimensional objects from all angles. Many animals and plant structures have a plane of bilateral symmetry. Here, we first (experiment I) give evidence that young poultry chicks readily generalize bilateral symmetry as a feature of two-dimensional patterns in fronto-parallel view. We then test the ability of chicks to recognize symmetry in images that would be produced by the transformed view produced by a 40° horizontal combined with a 20° vertical rotation of a pattern on a spherical surface. Experiment II gives evidence that chicks trained to distinguish symmetrical from asymmetrical patterns treat rotated views of symmetrical 'objects' as symmetrical. Experiment III gives evidence that chicks trained to discriminate rotated views of symmetrical 'objects' from asymmetrical patterns generalize to novel symmetrical objects either in fronto-parallel or rotated view. These findings emphasize the importance of bilateral symmetry for three-dimensional object recognition and raise questions about the underlying mechanisms of symmetry perception.     

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.     

Voltage-dependent channels permeable to K+ and Na+ in the membrane ofAcer pseudoplatanus vacuoles

Summary The patch-clamp technique in whole-cell configuration was used to study the electrical properties of the tonoplast in isolated vacuoles fromAcer pseudoplatanus cultured cells. In symmetrical KCl or K₂ malate solutions, voltage- and time-dependent inward currents were elicited by hyperpolarizing the tonoplast (inside negative), while in the positive range of potential the conductance was very small. The specific conductance of the tonoplast at –100 mV, in 100mm symmetrical KCl was about 160 S/cm². The reversal potentials (E _rev) of the current, measured in symmetrical or asymmetrical ion concentrations (cation, anion or both) were very close to the values of the K⁺ equilibrium potential. Experiments performed in symmetrical or asymmetrical NaCl indicate that Na⁺ too can flow through the channels. NeitherE _rev nor amplitude and kinetics of the current changed by replacing NaCl with KCl in the external solution. These results indicate the presence of hyperpolarization-activated channels in tonoplasts, which are permeable to K⁺ as well as to Na⁺. Anions such as Cl^– or malate seem to contribute little to the channel current.     

Locomotor structures in the Middle Triassic archosaurs from Los Chañares (La Rioja,Argentina)

A brief anatomical analysis of the pedial adaptations of the archosaurs from Los Chañares is made based on new available data.

The archosaurs from the Los Chañares local fauna show four different patterns in the structure of their hind limbs: (1) crurotarsal asymmetrical (Chanaresuchus and Tropidosuchus) (2) crurotarsal symmetrical (Gracilisuchus) (3) mesotarsal asymmetrical (Lagerpeton) (4) mesotarsal symmetrical (Lagosuchus). Each group has typical features that define “grades”; or morphological sets that were present at the same time in the local fauna, showing more derived characters than previously expected from the Middle Triassic.     

Walking at the preferred stride frequency maximizes local dynamic stability of knee motion

Healthy humans display a preference for walking at a stride frequency dependent on the inertial properties of their legs. Walking at preferred stride frequency (PSF) is predicted to maximize local dynamic stability, whereby sensitivity to intrinsic perturbations arising from natural variability inherent in biological motion is minimized. Previous studies testing this prediction have employed different variability measures, but none have directly quantified local dynamic stability by computing maximum finite-time Lyapunov exponent (λ^Max), which quantifies the rate of divergence of nearby trajectories in state space. Here, ten healthy adults walked 45 m overground while sagittal motion of both knees was recorded via electrogoniometers. An auditory metronome prescribed 7 different frequencies relative to each individual's PSF (PSF; ±5, ±10, ±15 strides/min). Stride frequencies were performed under both freely adopted speed (FS) and controlled speed (CS: set at the speed of PSF trials) conditions. Local dynamic stability was maximal (λ^Max was minimal) at the PSF, becoming less stable for higher and lower stride frequencies. This occurred under both FS and CS conditions, although controlling speed further reduced local dynamic stability at non-preferred stride frequencies. In contrast, measures of variability revealed effects of stride frequency and speed conditions that were distinct from λ^Max. In particular, movement regularity computed by approximate entropy (ApEn) increased for slower walking speeds, appearing to depend on speed rather than stride frequency. The cadence freely adopted by humans has the benefit of maximizing local dynamic stability, which can be interpreted as humans tuning to their resonant frequency of walking.     

Four lines of spermatid development and dimorphic spermatozoa in the sea urchin Anthocidaris crassispina (Echinodermata, Echinoida)

 The process of sperm development in the sea urchin Anthocidaris crassispina was studied by light and electron microscopy. Similar to other echinoids studied, a single flagellum, striated rootlet and nuage-like materials were present in spermatogonia of A. crassispina. Spermatocytes near the diplotene stage showed intracellular localization of the axoneme which appeared to be a retracted flagellum prior to cell division. Fibrous filaments were associated with a proximal centriole in spermatocytes and spermatids and might be involved in movement of the proximal centriole. An acrosomal vesicle was developed and a residual body was formed in spermatids. The special development patterns in A. crassispina attributed to the presence of two patterns of tail development and two patterns of mitochondrial development during spermiogenesis. These four lines of spermiogenesis resulted in the formation of four morphological types of sperm cell, i.e. sperms with: (1) a symmetrical midpiece and posterior tail, (2) an asymmetrical midpiece and posterior tail, (3) a symmetrical midpiece and bent tail and (4) an asymmetrical midpiece and bent tail. Sperm cells with bent tails (type 3+4) were probably still at the late spermatid stage because results of scanning electron microscopy demonstrated gradual detachment and eventual straightening of the bent tail, and their percentage occurrence in the sperm population decreased significantly (P<0.05) towards the spawning season of A. crassispina. Spermatozoa with a symmetrical midpiece were dominant (averaging 70% occurrence in the sperm population) over those with an asymmetrical midpiece. The dimorphic spermatozoa in A. crassispina (types 1, 2) are both considered to be euspermatozoa as their morphology is typical for Echinoida. Accepted: 4 May 1998     

Locomotion energetics and gait characteristics of a rat-kangaroo,<Emphasis Type="Italic"> Bettongia penicillata</Emphasis>, have some kangaroo-like features

The locomotory characteristics of kangaroos and wallabies are unusual, with both energetic costs and gait parameters differing from those of quadrupedal running mammals. The kangaroos and wallabies have an evolutionary history of only around 5 million years; their closest relatives, the rat-kangaroos, have a fossil record of more than 26 million years. We examined the locomotory characteristics of a rat-kangaroo, Bettongia penicillata. Locomotory energetics and gait parameters were obtained from animals exercising on a motorised treadmill at speeds from 0.6 m s⁻¹ to 6.2 m s⁻¹. Aerobic metabolic costs increased as hopping speed increased, but were significantly different from the costs for a running quadruped; at the fastest speed, the cost of hopping was 50% of the cost of running. Therefore B. penicillata can travel much faster than quadrupedal runners at similar levels of aerobic output. The maximum aerobic output of B. penicillata was 17 times its basal metabolism. Increases in speed during hopping were achieved through increases in stride length, with stride frequency remaining constant. We suggest that these unusual locomotory characteristics are a conservative feature among the hopping marsupials, with an evolutionary history of 20–30 million years. Communicated by I.D. Hume An erratum to this article can be found at     

Distal Forelimb Kinematics in <Emphasis Type="Italic">Erythrocebus patas</Emphasis> and <Emphasis Type="Italic">Papio anubis</Emphasis> During Walking and Galloping

When using symmetrical gaits, terrestrial digitigrade monkeys adopt less digitigrade, i.e., more palmigrade-like, hand postures as they move with faster speeds. Accordingly, it appears that, in contrast to other mammals, digitigrady is unrelated to cursoriality in primates. However, researchers have not documented the effects of speed on distal forelimb kinematics in faster asymmetrical gaits, i.e., galloping, when ground reaction forces are typically increased owing to the decreased number of contact points during a stride, combined with higher speed. Thus, it remains possible that primates use digitigrade hand postures during these higher-speed asymmetrical gaits. We investigated 3D angles in the wrist joint and metacarpophalangeal joint of 2 habitually digitigrade terrestrial monkeys, Erythrocebus patas and Papio anubis, across a large range of walking and galloping speeds on a motorized treadmill. Nonparametric analyses reveal that angles, and therefore hand postures, are not different at the subject’s walk-gallop transition. Regression analyses show that when walking, digitigrade postures are adopted at slow speeds and more palmigrade-like postures are adopted at fast speeds. Contrary to expectations, there is little change in hand postures across galloping speeds; both subjects maintained palmigrade-like hand postures with substantial joint yield and reextension during support. These results indicate that the hands are always less digitigrade at faster speeds because the joints of the distal forelimb cannot resist the higher ground reaction forces that accompany these higher speed gaits.     

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.     

Dynamics of the Terrestrial Mammals of Ipolytarnóc (Northern Hungary)

Much has been discovered about the movement speed of extant animals through the use of a variety of instruments, but much less is known about the speeds of extinct animals. The anatomy and mobility of prehistoric animals can be deduced from their bone remains and ichnofossils. By the use of biomechanics, their movement speeds also can be determined. Many ichnofossils have been found at Ipolytarnóc (the Miocene footprint locality at Ipolytarnóc, Hungary.) Ichnofossils of the most studied mammals from this site have been used in this study: the herbivores Rhinoceripeda tasnadyi, Megapecoripeda miocaenica, and Pecoripeda hamori, and the carnivore Mustelipeda punctata. The movement speed of each species was calculated from its foot and stride lengths. At Ipolytarnóc, Rhinoceripeda tasnadyi was able to move with a speed of 4.29 m/s, while Megapecoripeda miocaenica could reach a speed of 3.01 m/s. In the case of Pecoripeda hamori, its speed was 4.53 m/s, while in the case of Mustelipeda punctata it was 2.05 m/s. Conclusions about the paleoenvironment and the movement speeds of the prehistoric animals are based on our results. At times, the ancient mammals of Ipolytarnóc wallowed in puddles and muddy pools.     

Lithium treatment of Nicotiana tabacum microspores blocks polar nuclear migration,disrupts the partitioning of membrane-associated Ca2+, and induces symmetrical mitosis

We have found that the normal developmental pathway of Nicotiana tabacum microspores is blocked or switched when microspores are exposed to lithium, and these effects are reversible with Ca²⁺ and myo-inositol. Normal development was defined by the following characteristics: changes in microspore shape from spherical to oval and then ellipsoid; two nuclear displacements, first from a central location to the cell periphery, and then from the periphery to the generative pole; a localization of membrane-associated Ca²⁺ at the generative pole preceding nuclear division; and, finally, an asymmetrical mitosis that results in a two-celled pollen grain with well-differentiated generative and vegetative nuclei. Lithium treatment blocked the localization of membrane-associated Ca²⁺ at the generative pole, and instead it was evenly distributed at both poles. Lithium treatment also blocked the asymmetrical positioning of the microspore nucleus at the generative pole and resulted in an approximately four-fold increase in the frequency of symmetrical mitosis. When Ca²⁺ and myo-inositol were added along with lithium, the effects were substantially decreased, and there was only a small increase in the frequency of symmetrical mitosis compared with controls. The timing of treatment was important; microspores isolated before the first nuclear displacement had a low frequency of further development, while microspores isolated immediately preceding the onset of mitosis were much less sensitive to lithium, and the result was only a small increase in the frequency of symmetrical mitosis. In microspores isolated after the first nuclear displacement, a 1-day exposure to lithium was sufficient to switch the developmental pathway from an asymmetrical to a symmetrical mitosis while still allowing limited further development. However, we have not optimized culturing conditions for embryogenesis and the furthest development observed after a 1-week culture was to four- or five-celled proembryo-like structures.     

Effect of nitrogen and phosphorus starvation on the polyunsaturated triacylglycerol composition, including positional isomer distribution, in the alga Trachydiscus minutus

The yellow-green alga Trachydiscus minutus (Eustigmatophyceae, Heterocontophyta) was cultivated in a standard medium and under nitrogen- and phosphorus-starvation and its triacylglycerols were analyzed by RP-HPLC/MS-APCI. The molecular species of triacylglycerols included a total of 74 triacylglycerols having at least one polyunsaturated fatty acid. Polyunsaturated triacylglycerols were identified for the first time in a yellow-green alga. N-starvation brought about a nearly 50% drop in TAGs containing EPA, and also decreased TAGs containing ARA, while P-starvation had a sizable effect on those TAGs that contain two or three arachidonic acids. In four TAGs containing PUFA, i.e. EEE, EEA, EAA and AAA, N-starvation caused a rapid fivefold increase in ARA content and the ratio of TAGs containing ARA, i.e. AEE to AAA increased tenfold relative to control. Regioisomeric characterization of triacylglycerols containing palmitic, arachidonic (ARA) and eicosapentaenoic acids (EPA) showed that the proportion of positional isomers is affected by N- and P-starvation. N- and P-starvation also changed the ratio of symmetrical to asymmetrical TAGs. Positional isomers exhibited identical ratios of symmetrical and asymmetrical TAGs irrespective of the type of FAs. In control cultivation the major TAGs with a single PUFA were symmetrical ones (PEP or PAP) whose ratio to asymmetrical counterparts (PPE or PPA) was about 3:1, whereas N- and P-starvation yielded opposite ratios, 1:3–1:5. The control cultivation yielded ∼90% asymmetrical TAGs with two PUFAs (i.e. PEE and PAA), whereas with N- and P-starvation the ratio of symmetrical to asymmetrical TAGs increased to 2:1 and 3:2, respectively.     

Percutaneous osseointegrated prostheses for amputees: Limb compensation in a 12-month ovine model

Percutaneous osseointegrated prostheses are being investigated as an alternative strategy to attach prosthetic limbs to patients. Although the use of these implants has shown to be promising in clinical trials, the ability to maintain a skin seal around an osseointegrated implant interface is a major challenge to prevent superficial and deep periprosthetic infections. The specific aim of this study was to establish a translational load-bearing ovine model to assess postoperative limb compensation and gait symmetry following a percutaneous osseointegrated implant. We tested the following hypotheses: (1) the animals would return to pre-amputation limb loads within 12-months; (2) the animals would return to a symmetrical gait pattern (stride length and time in stance) within 12-months. The results demonstrated that one month following surgery, the sheep loaded their amputated limb to a mean value of nearly 80% of their pre-amputation loading condition; by 12-months, this mean had dropped to approximately 74%. There was no statistical differences between the symmetry of the amputated forelimb and the contralateral forelimb at any time point for the animals stride length or the time spent in the stance phase of their gait cycle. Thus, the data showed that while the animals maintained symmetric gait patterns, they did not return to full weight-bearing after 12-months. The results of this study showed that a large animal load-bearing model had a symmetric gait and was weight bearing for up to 12 months. While the current investigation utilizes an ovine model, the data show that osseointegrated implant technology with postoperative follow-up can help our human patients return to symmetric gait and maintain an active lifestyle, leading to an improvement in their quality of life following amputation.     

The effects of natural substrate discontinuities on the quadrupedal gait kinematics of free‐ranging Saimiri sciureus

Wild primates encounter complex matrices of substrates that differ in size, orientation, height, and compliance, and often move on multiple, discontinuous substrates within a single bout of locomotion. Our current understanding of primate gait is limited by artificial laboratory settings in which primate quadrupedal gait has primarily been studied. This study analyzes wild Saimiri sciureus (common squirrel monkey) gait on discontinuous substrates to capture the realistic effects of the complex arboreal habitat on walking kinematics. We collected high‐speed video footage at Tiputini Biodiversity Station, Ecuador between August and October 2017. Overall, the squirrel monkeys used more asymmetrical walking gaits than symmetrical gaits, and specifically asymmetrical lateral sequence walking gaits when moving across discontinuous substrates. When individuals used symmetrical gaits, they used diagonal sequence gaits more than lateral sequence gaits. In addition, individuals were more likely to change their footfall sequence during strides on discontinuous substrates. Squirrel monkeys increased the time lag between touchdowns both of ipsilaterally paired limbs (pair lag) and of the paired forelimbs (forelimb lag) when walking across discontinuous substrates compared to continuous substrates. Results indicate that gait flexibility and the ability to alter footfall patterns during quadrupedal walking may be critical for primates to safely move in their complex arboreal habitats. Notably, wild squirrel monkey quadrupedalism is diverse and flexible with high proportions of asymmetrical walking. Studying kinematics in the wild is critical for understanding the complexity of primate quadrupedalism.     

A new way of analysing symmetrical and asymmetrical gaits in quadrupeds

This new approach highlights for the first time that all the gaits, symmetrical as well as asymmetrical, correspond to the succession of sequences that start by the movements of the two fore limbs, followed by the movements of the two hind limbs. Inside those sequences, the gaits are identified by the time lag between the movements of the two pairs and by the time lag between the movements of the two feet inside each pair. This approach, by breaking the stride paradigm, gives a new framework to locomotion analysis.     

Effects of growth and speed on hindlimb joint angular displacement patterns in vervet monkeys (Cercopithecus aethiops)

Hip, knee, and ankle joint displacement patterns are compared across both age and speed for five immature vervet monkeys sampled approximately every 6 months over a 3 year period. The analysis indicated that, as a group, the animals displayed no consistent changes in joint patterns as they grew. However, individual animals showed consistent patterns. There were also no consistent effects of size across animals at the walk-gallop transition. This is contrary to McMahon's prediction (J. Appl. Physiol. 39:619-627, 1975) based upon his elastic-similarity model of animal scaling. With increasing speed, when symmetrical gaits were used, all of the animals tended to show a decrease in the relative positions of the hip, knee, and ankle maximum values. Furthermore, across the walk-gallop transition, the animals tended to show a decrease in the range of ankle and knee movements.     

Random segregation inNeurospora

Data from five-point crosses involving linkage group I and three-point crosses involving linkage groups VI and VII ofNeurospora crassa andN. sitophila have been analysed in order to detect the phenomena of polarized segregation and partial spindle overlap.The results clearly show absence of polarized segregation in both species. They further show that partial spindle overlap is a rare event if at all it occurs in the two species. The occurrence of a majority of either symmetrical or asymmetrical classes is a feature common to both species.     

Karyotypic analysis of Cajanus,Atylosia, and Rhynchosia species

Summary Somatic chromosomes of two cultivais of Cajanus cajan, eight species of Atylosia (A. albicans, A. cajanifolia, A. lineata, A. platycarpa, A. scarabaeoides, A. serica, A. trinervia and A. volubilis), and of Rhynchosia rothii were analysed. All species had 2n=22. Eight of the 10 species studied had two pairs of satchromosomes while A. scarabaeoides and A. sericea had only one sat-chromosome pair. Based on relative chromosome length (L%), arm ratio (pa-value) and presence or absence of secondary constriction, a karyotype formula for each species was formulated. Based on these parameters the chromosome pairs could also be assigned to groups ranging from 8 to 10 in different species. Except for the asymmetrical karyotype of A. albicans, the other species had rather moderately symmetrical karyotypes.