The gait of children with and without cerebral palsy: work, energy, and angular momentum

This paper describes a method to characterize gait pathologies like cerebral palsy using work, energy, and angular momentum. For a group of 24 children, 16 with spastic diplegic cerebral palsy and 8 typically developed, kinematic data were collected at the subjects self selected comfortable walking speed. From the kinematics, the work-internal, external, and whole body; energy-rotational and relative linear; and the angular momentum were calculated. Our findings suggest that internal work represents 53% and 40% respectively of the whole body work in gait for typically developed children and children with cerebral palsy. Analysis of the angular momentum of the whole body, and other subgroupings of body segments, revealed a relationship between increased angular momentum and increased internal work. This relationship allows one to use angular momentum to assist in determining the kinetics and kinematics of gait which contribute to increased internal work. Thus offering insight to interventions which can be applied to increase the efficiency of bipedal locomotion, by reducing internal work which has no direct contribution to center of mass motion, in both normal and pathologic populations.     

Patterns and processes in the early evolution of the tetrapod ear

This article reviews some of the latest information on the evolution of the tetrapod ear region as seen in the fossil record. It looks at the changes that can be documented across the fish-tetrapod transition, the patterns that they show and what can be inferred of the processes that brought some of them about. These processes include an increased role for neural crest, and heterochronic processes such as pedomorphosis. The earliest tetrapods show a common pattern of a short stout stapes with a large stapedial foramen, that primitively contacted the palatal bones and probably supported the braincase. Modifications to this pattern can be seen in tandem with changes to the occiput and are bound up with changes to jaw and breathing mechanisms. By the Late Carboniferous, tetrapods had diversified into a range of groups showing a wide variety of otic morphologies, some of which were probably tympanic, while others were not, and some which are very different from those found in extant tetrapods. In amniotes, the evolution of a tympanic ear appears to correlate with consolidation and integration of the occiput to the skull roof. Competing phylogenies suggest different numbers of iterations for the origin of a tympanic ear, but a minimum of four separate occasions is implied.     

Devonian climate change, breathing, and the origin of the tetrapod stem group

The diversification of the tetrapod stem group occurred duringthe late Middle through the Late Devonian, that is from theGivetian to Famennian stages about 385–365 million yearsago. The relationships between the known taxa representing thisradiation have currently reached a reasonable consensus so thatinterpretations of the order of appearance of tetrapod charactersis possible. The immediate fish relatives of the earliest limbedtetrapods show what is interpreted as a progressive increasein the spiracular chamber and its opening to the outside. Here,this is inferred to be associated with an increased capacityfor air-breathing. Lungs are thought to have been present inmost early bony fishes, and were most likely ventilated by air-gulping.This could have brought about a facultative capacity for air-breathing,which the tetrapod stem group exploited to the greatest degree.These adaptations are shown not only in freshwater forms butalso in estuarine and marginal marine forms. Estimates of oxygenlevels during this period suggest that they were unprecedentedlylow during the Givetian and Frasnian periods. At the same time,plant diversification was at its most rapid, changing the characterof the landscape and contributing, via soils, soluble nutrients,and decaying plant matter, to anoxia in all water systems. Theco-occurrence of these global events may explain the evolutionof air-breathing adaptations in at least two lobe-finned groups,contributing directly to the rise of the tetrapod stem group.In contrast to recent studies, low atmospheric oxygen is notconsidered to be a causal factor in the lack of fossils documentingthe evolution of Early Carboniferous tetrapods.     

A comparison of muscular mechanical energy expenditure and internal work in cycling

The hypothesis that the sum of the absolute changes in mechanical energy (internal work) is correlated with the muscular mechanical energy expenditure (MMEE) was tested using two elliptical chainrings, one that reduced and one that increased the internal work (compared to circular). Upper and lower bounds were put on the extra MMEE (work done by net joint torques in excess of the external work) with respect to the effect of intercompensation between joint torques due to biarticular muscles. This was done by having two measures of MMEE, one that allowed no intercompensation and one that allowed complete intercompensation between joints spanned by biarticular muscles. Energy analysis showed no correlation between internal work and the two measures of MMEE. When compared to circular, the chainring that reduced internal work increased MMEE, and phases of increased crank velocity associated with the elliptical shape resulted in increased power absorbed by the upstroke leg as it was accelerated against gravity. The resulting negative work necessitated additional positive work. Thus, the hypothesis that the internal work is correlated with MMEE was found to be invalid, and the total mechanical work done cannot be estimated by summing the internal and external work. Changes in the dynamics of cycling caused by a non-circular chainring may affect performance and must be considered during the non-circular chainring design process.     

First report of rapid eye color change in a non-avian tetrapod

Like coloration of the integument, eye color can be a significant but understudied component of communication and reproductive behavior. Eye color can change with sexual maturation and become sexually dimorphic, but in a few birds and fish, eye color can also change rapidly in response to the environment. There are few cases of the latter, and we report here several instances of such change in eye color in the Eastern box turtle (Terrapene carolina carolina), the first non-avian tetrapod in which this capability has been reported. In male turtles, the iris changed from a pale yellow color (often characteristic of juveniles) to a bright red color (characteristic of mature males) in a period of <5 s. The nature of the color change is similar to that observed in some birds and suggests a common mechanism and/or adaptive role, which could be further explored in Eastern box turtles.     

Patterns of bryophyte and vascular plant richness in European subalpine springs

The diversity of spring habitats can be determined not only by local environmental conditions, but also by large-scale biogeographical effects. The effects can differ across various groups of organisms. We compared α-, β- and γ-diversity patterns of bryophytes and vascular plants of (sub)alpine springs in three contrasting mountain ranges: Alps (Switzerland), Balkans (Bulgaria), Western Carpathians (Slovakia, Poland). We used univariate and multivariate statistics to test for the effects of pH, conductivity, altitude, slope, mean annual temperature and annual precipitation on diversity patterns of both taxonomic groups and compared diversity patterns among the regions for particular pH and conductivity classes. We identified acidophyte and basiphyte, calcifuge and calcicole species using species response modelling. All regions displayed significant relationship between conductivity and α-diversity of vascular plants. Bulgaria showed the highest α-diversity of vascular plants for the middle part of the conductivity gradient. For both taxonomic groups, the β-diversity in the middle part of gradient was highest in Swiss Alps. The total species pool was lowest in Bulgaria. The percentage of basiphyte and calcicole species was highest in the Alps. In (sub)alpine springs, mineral richness was a better determinant of vascular plant α-diversity than pH, and the extent of the alpine area did not coincide with α-diversity. Observed inter-regional differences in diversity patterns could be explained by the different proportion of limestone bedrock and different biogeographic history. The differences in α-diversity between both taxonomic groups are presumably result of the different rates of adaptation processes.     

Developmental mechanisms of macroevolutionary change in the tetrapod axis: A case study of Sauropterygia

Understanding how developmental processes change on macroevolutionary timescales to generate body plan disparity is fundamental to the study of vertebrate evolution. Adult morphology of the vertebral column directly reflects the mechanisms that generate vertebral counts (somitogenesis) and their regionalisation (homeotic effects) during embryonic development. Sauropterygians were a group of Mesozoic marine reptiles that exhibited an extremely high disparity of presacral vertebral/somite counts. Using phylogenetic comparative methods, we demonstrate that somitogenesis and homeotic effects evolved in a co‐ordinated way among sauropterygians, contrasting with the wider pattern in tetrapods, in which somitogenetic and homeotic shifts are uncorrelated. Changes in sauropterygian body proportions were primarily enabled by homeotic shifts, with a lesser, but important, contribution from differences in postpatterning growth among somites. High body plan plasticity was present in Triassic sauropterygians and was maintained among their Jurassic and Cretaceous descendants. The extreme disparity in the body plan of plesiosaurian sauropterygians did not result from accelerated rates of evolutionary change in neck length, but instead reflect this ancestral versatility of sauropterygian axial development. Our results highlight variation in modes of axial development among tetrapods, and show that heterogeneous statistical models can uncover novel macroevolutionary patterns for animal body plans and the developmental mechanisms that control them.     

Fundamental energetics of birds: 1. The maximum ability of birds to change their thermal conductance and the efficiency of metabolic energy transformation into mechanical work

The maximum abilities of birds to dissipate heat with an increase in their metabolic rate as a result of both activity and heat stress have been estimated by an integrated measurement of energy expenditure at rest (measured according to oxygen consumption) and common activity, the so-called existence metabolism (measured according to food consumption) in 26 passerine species, covering the overall order relative to the size range (from goldcrest to raven) and 16 nonpasserine species of the same size range (25?C4000 g). The passerine and nonpasserine species do not differ in the mean change of heat loss in the cases of both maximal (h _min) and minimal (h _max) insulations. Both the minimal (h _min) and maximal (h _max) nonevaporative heat losses in both groups of species display similar dependences on the body weight, and the regression slopes for both h _min and h _max are equal. On the other hand, h _max is approximately fourfold higher as compared with h _min. This demonstrates that birds at the same ambient temperature (T _A) are able to increase fourfold the amount of dissipated heat losses without increasing their evaporative heat loss. An increase in the h _max/h _min ratio suggests a more perfect organization of the systems associated with blood circulation and respiration, which naturally gives great advantages for any life activities, first and foremost, allowing for an increase in activity. The efficiency of metabolic energy transformation into mechanical work during standard existence varies in the studied species from 0.28 to 0.22 in summer and in winter.     

Joit torque and energy patterns in normal gait

Experiments were conducted on normal level gait to determine the synergistic patterns present in the forces causing joint moments and those associated with the generation, absorption and transfer of mechanical energy. The following generalizations can be made about the patterns: (i) During swing phase three forces (gravitational, muscle and knee joint acceleration) are responsible for shank rotation, and are shown to act together during both acceleration and deceleration.—(ii) The patterns of generation, absorption and transfer of mechanical energy at the joints are detailed. These patterns demonstrate inter-segment transfers of energy through the joint centres, and through the muscles, as well as the more recognized generation and absorption by the muscles themselves.—As a result of the complexity shown in these patterns it is cautioned that fundamental relationships that may have been derived from controlled biomechanical experiments (such as horizontal flexion and extension of the forearm) are not likely to apply to more normal movements such as gait.     

Determination of equivalent amounts of kinetic energy, work, and heat energy in the human body

The goal of this study is determine the mechanical equivalent of heat and the functional capacity of metabolism of walking at a slow pace (velocity = 4022m/hour, length of a step=75cm, energy utilization of a 70 kg person is 200kcal/hour). 50 healthy physicians were chosen randomly, and up and down motion of the body were determined as 6cm while stepping. Based on these, the heat equivalent is 37.5kcal/hour for horizontal motion and 52.7kcal/hour for 6cm up-and-down bobbing motions of body, and the functional capacity of metabolism is at least 45% ([37.5+52.7]/200=45%) for slow walking state, that this capacity is twofold more than earlier information. Muscle converts kinetic energy (work) to heat via friction, and heat sources of the body, and the concepts of thermogenesis and the functional capacity of metabolism should be revised.     

Getting a grip on tetrapod grasping: form,function, and evolution

Human beings have been credited with unparalleled capabilities for digital prehension grasping. However, grasping behaviour is widespread among tetrapods. The propensity to grasp, and the anatomical characteristics that underlie it, appear in all of the major groups of tetrapods with the possible exception of terrestrial turtles. Although some features are synapomorphic to the tetrapod clade, such as well‐defined digits and digital musculature, other features, such as opposable digits and tendon configurations, appear to have evolved independently in many lineages. Here we examine the incidence, functional morphology, and evolution of grasping across four major tetrapod clades. Our review suggests that the ability to grasp with the manus and pes is considerably more widespread, and ecologically and evolutionarily important, than previously thought. The morphological bases and ecological factors that govern grasping abilities may differ among tetrapods, yet the selective forces shaping them are likely similar. We suggest that further investigation into grasping form and function within and among these clades may expose a greater role for grasping ability in the evolutionary success of many tetrapod lineages.     

Patterns of vegetation surrounding springs in Goshen Bay,Utah County,Utah U.S.A.

Saline meadows in Goshen Bay, Utah County, Utah, USA, occur interspersed among large areas of salt playa. Springs irrigate these meadows which show concentric vegetational zones surrounding each spring. Each vegetational zone demonstrates its own unique dominant vascular plant species. The central zone is dominated primarily by Scirpus americanus, the middle zone by Eleocharis palustris and the outer zone by Juncus balticus, Distichlis spicata and Muhlenbergia asperifolia. As distance increases from the springs, pH and soluble salt concentration increase, while percent organic matter, percent moisture and phosphorus decrease. With the increase in salt levels and decrease in water levels, halophytic plant species generally increase with distance from the springs.     

Old men running: mechanical work and elastic bounce

It is known that muscular force is reduced in old age. We investigate what are the effects of this phenomenon on the mechanics of running. We hypothesized that the deficit in force would result in a lower push, causing reduced amplitude of the vertical oscillation, with smaller elastic energy storage and increased step frequency. To test this hypothesis, we measured the mechanical energy of the centre of mass of the body during running in old and young subjects. The amplitude of the oscillation is indeed reduced in the old subjects, resulting in an approximately 20% smaller elastic recovery and a greater step frequency (3.7 versus 2.8 Hz, p=1.9x10(-5), at 15-17 km h(-1)). Interestingly, the greater step frequency is due to a lower aerial time, and not to a greater natural frequency of the system, which is similar in old and young subjects (3.6 versus 3.4 Hz, p=0.2). Moreover, we find that in the old subjects, the step frequency is always similar to the natural frequency, even at the highest speeds. This is at variance with young subjects who adopt a step frequency lower than the natural frequency at high speeds, to contain the aerobic energy expenditure. Finally, the external work to maintain the motion of the centre of mass is reduced in the old subjects (0.9 versus 1.2 J kg(-1) m(-1), p=5.1x10(-6)) due to the lower work done against gravity, but the higher step frequency involves a greater internal work to reset the limbs at each step. The net result is that the total work increases with speed more steeply in the old subjects than in young subjects.     

Amplitude effects of medio-lateral mechanical and visual perturbations on gait

Falls during walking are a major contributor to accidental deaths and injuries that can result in debilitating hospitalization costs, lost productivity, and diminished quality of life. To reduce these losses, we must develop a more profound understanding of the characteristic responses to perturbations similar to those encountered in daily life. This study addresses this issue by building on our earlier studies that examined mechanical and visual perturbations in the same environment by applying the same continuous pseudo-random perturbations at multiple (3 mechanical, 5 visual) amplitudes. Walking variability during mechanical perturbations increased significantly with amplitude for all subjects and differences as measured by variabilities of step width, COM position, and COM velocity. These parameters were the only ones sensitive to the presence of visual perturbations, but none of them changed significantly with perturbation amplitude. Additionally, visual perturbation effects were far less consistent across participants, with several who were essentially unaffected by visual perturbations at any level. The homogeneity of the mechanical perturbation effects demonstrates that human responses to mechanical perturbations are similar because they are driven by kinetics that require similar corrections that must be made in order to maintain balance. Conversely, responses to visual perturbations are driven by the perceived need to make corrections and this perception is not accurate enough to produce amplitude-related corrections, even for a single participant, nor is this perception consistent across individuals. This latter finding is likely to be relevant to future visual perturbation studies and the diagnosis and rehabilitation of gait and balance disorders.     

Cretaceous small scavengers: feeding traces in tetrapod bones from Patagonia, Argentina

Ecological relationships among fossil vertebrate groups are interpreted based on evidence of modification features and paleopathologies on fossil bones. Here we describe an ichnological assemblage composed of trace fossils on reptile bones, mainly sphenodontids, crocodyliforms and maniraptoran theropods. They all come from La Buitrera, an early Late Cretaceous locality in the Candeleros Formation of northwestern Patagonia, Argentina. This locality is significant because of the abundance of small to medium-sized vertebrates. The abundant ichnological record includes traces on bones, most of them attributable to tetrapods. These latter traces include tooth marks that provde evidence of feeding activities made during the sub-aerial exposure of tetrapod carcasses. Other traces are attributable to arthropods or roots. The totality of evidence provides an uncommon insight into paleoecological aspects of a Late Cretaceous southern ecosystem.     

Comparable disparity in the appendicular skeleton across the fish–tetrapod transition,and the morphological gap between fish and tetrapod postcrania

Appendicular skeletal traits are used to quantify changes in morphological disparity and morphospace occupation across the fish–tetrapod transition and to explore the informativeness of different data partitions in phylogeny reconstruction. Anterior appendicular data yield trees that differ little from those built from the full character set, whilst posterior appendicular data result in considerable loss of phylogenetic resolution and tree branch rearrangements. Overall, there is a significant incongruence in the signals associated with pectoral and pelvic data. The appendicular skeletons of fish and tetrapods attain similar levels of morphological disparity (at least when data are rarefied at the maximum sample size for fish in our study) and occupy similarly sized regions of morphospace. However, fish appear more dispersed in morphospace than tetrapods do. All taxa show a heterogeneous distribution in morphospace, and there is a clear separation between fish and tetrapods despite the presence of several evolutionarily intermediate taxa.