Morphological changes leading to hominid bipedalism

The adoption of habitual bipedal locomotion required a backward shift of the centre of gravity of the body, to a level relative to the supporting surface area of the body and pivotal axis of the hips at which walking with extended knees became practicable. In the morphology of the immediate ancestors to hominids, there were relatively few features whose change could have effectively affected the position of the gravity vertical of the body. Weight could be distributed posteriorly mainly by increasing the mass of the hindlimbs and the kyphotic curve of the vertebral column, by developing buttocks. and by flattening the thorax and abdomen. Estimates indicate that the adoption of a suitable curvature of the vertebral column alone was not sufficient for shifting the centre of gravity behind the 'threshold of bipedalism'. A considerable increase in the mass of the hindlimbs was also required, and the addition of the mass of the buttocks may have represented the decisive factor for crossing the threshold. A possible physical enviroment in which the change to bipedalism could have taken place was a mountainous terrain with long, steep slopes, transected by gorges and precipices. In such a terrain the increase in the power and mass of both the hindlimbs and the gluteus maximus proprius muscle could have been favoured by selection, leading ultimately to a condition in which the combined effect of heavy hindlimbs. a suitable curvature of the spine and the weight of the buttocks shifted the centre of gravity of the body backwards to a level at which habitual walking with extended knees became practicable. □ Hominid evolution. bipedalism, evolutionary thresholds.     

Proportions and function of the limbs of glyptodonts

Vizcaíno, S.F., Blanco, R.E., Bender, J.B. & Milne, N. 2010: Proportions and function of the limbs of glyptodonts. Lethaia, Vol. 44, pp. 93–101. This study examines the limb bone proportions and strength of glyptodonts (Xenarthra, Cingulata). Two methods are used to estimate the body mass and location of the centre of gravity of the articulated specimens. These estimates, together with measurements of the femur and humerus, are used to calculate strength indicators (SI). The other long bones of the limbs are used to calculate limb proportion indices that give an indication of digging ability, speed, and limb dominance in armadillos, the glyptodonts’ living closest relatives. The results show that regardless of how the body mass and centre of gravity are calculated, the majority of the glyptodont’s weight is borne by the hindlimbs. The SI calculations show that femora are sturdy enough to bear these loads. The fact that the femora have higher SI than the humerii indicates that sometimes the hindlimbs are required to bear an even greater proportion of the body weight, possibly when rising to a bipedal posture or pivoting on their hindlimbs to deliver a blow with their armoured tail. The analysis of limb proportions indicates that both the hindlimb and the forelimb have proportions that correlate strongly with body mass. This outcome supports the other results, but also shows that forelimbs must be also involved in manoeuvring the glyptodont body. □Glyptodonts, Mammalia, Xenarthra, limbs, strength indicators.     

OBSERVATIONS ON THE DIETARY CHOICE OF FREE-RANGING JUVENILE OSTRICHES

Cooper, S.M. & Palmer, T. 1994. Observations on the dietary choice of free-ranging juvenile Ostriches. Ostrich 65:251-255.

The dietary selection of juvenile Ostriches in an east African savanna was determined using penned and free-ranging chicks. The young Ostriches displayed an innate preference for selected plant species and showed little change free-ranging chicks. The young Ostriches displayed increasing experience. The most heavily utilized foods were newly emergent grass leaves and the foliage of the woody forbs Indigofera schimperi (Leguminosae) and Aspilia mossambicensis (Comositae) While the diet composition of the Ostrich is similar to that of Grant's gazelle, Ostriches differ in that they eat some silica rich plants not utilized by ungulates; thus Ostriches may be a useful addition to mixed-species game ranching in Africa.     

The legs of ostriches (Struthio) and moas (Pachyornis)

Ostriches were filmed running at maximum speed, and forces on the feet were calculated. Measurements were made of the principal structures in the legs of an ostrich. Hence peak stresses in muscles, tendons and bones were calculated. They lay within the range of stresses calculated for strenuous activities of other vertebrates. The ostrich makes substantial savings of energy in running, by elastic storage in stretched tendons. Pachyornis was a flightless bird, much heavier than ostriches and with massively thick leg bones. These bones are shorter than predicted for its estimated body mass, by extrapolation from allometric equations for flying birds. An attempt is made to calculate the stresses that acted in the leg bones in running, for all possible patterns of leg movement. The stresses were probably rather low, unless Pachyornis was capable of running fast. It is argued that the optimum factor of safety for moo leg bones may have been exceptionally high, as a consequence of the absence of predators.     

The functions of the lumbar spine during stepping in the cat

To examine the functional roles played by the lumbar spine during overground stepping, seven adult cats were run in electromyographic (EMG) experiments. Recordings were made bilaterally from mm. iliocostalis, longissimus dorsi and multifidus at a single vertebral level (L₃) and from m. rectus abdominis. Stepping movements were monitored synchronously either by videotape or by high speed cinematography. During alternate use of the hindlimbs (walking and trotting), both epaxial and abdominal muscles were active bilaterally and biphasically. During in-phase use of the hindlimbs (galloping and half-bounding), single bursts of activity were observed. Phasic bursts of activity in rectus abdominus were reciprocal to those of epaxial muscles. Second bursts of activity in either group were noted infrequently. Recordings from the same back muscle at several vertebral levels indicated little difference from these patterns. Movements of the lumbar spine during galloping and half-bounding steps, both angular and linear, are easily correlated with muscle activity patterns. Movements of the lumbar spine during walking and trotting show no particular pattern. Only small angular and linear movements are found. It is concluded that the lumbar spine contributes substantially to step length and limb speed during galloping and half-bounding steps and the epaxial and abdominal musculature may also act as elastic bodies. During walking and trotting steps, the epaxial muscles are proposed to act to stabilize the pelvic girdle to provide a firm base for limb muscles which arise on the pelvis and are synchronously active.     

4. CLIMPSES OF THE NARINA TROGON

Bertram, B. C. R. &; Burger, A. E. 1981. Aspects of incubation in Ostriches. Ostrich 52:36-43.

We studied incubation in domesticated Ostriches Struthio camelus in South Africa and wild Ostriches in Kenya. Although the eggs were large, with relatively high thermal capacities, unattended eggs exposed to the sun reached dangerously high temperatures (40,5°C). Experimental exposure of fresh eggs to the sun for seven days prior to incubation greatly reduced the percentage of embryos which developed, and no embryos survived 15 days of exposure. In the wild. Ostriches frequently shade their eggs in the pre-incubation period to prevent overheating.

During natural incubation, temperatures in the eggs (range 30,8-33,8°C) and of nest-air (31,9-34,6°C) were remarkably constant, despite the daily ambient fluctuations of air temperatures (17,8-38,9°C). Similarly the humidity of the nest-air (39–52%) was lower and less variable than the ambient air (39–72%). Water loss during 42 days of incubation was 11–12% of initial egg weight and, in addition, early laid eggs lost 3–4% during the 2½-3 week pre-incubation period. The water vapour conductivity and the daily water loss of Ostrich eggs were similar to those of other birds, in proportion to epg size, despite the arid environment inhabited by Ostriches. Some of the constraints on the feeding and breeding behaviour of Ostriches imposed by the physical requirements of their eggs are discussed.     

Mechanics of running of the ostrich (Struthio camelus)

Ostriches have been filmed running fast in their natural habitat. A female ostrich has been dissected and the principal bones, muscles and tendons in a leg have been measured. It is calculated that stresses up to 240 kN m⁻² and 40 MN m⁻², respectively, act in the digital flexor muscles and their tendons during running. Tensile and compressive stresses up to about 70MNm⁻² and 110 MNm⁻² act in the tibiotarsus. A large proportion of the energy which would otherwise be required for running is probably saved by elastic storage in tendons. Comparisons are made with the legs of flying birds and of antelopes.     

Brachially innervated ectopic hindlimbs in the chick embryo. I. Limb motility and motor system anatomy during the development of embryonic behavior

The functional status of brachially innervated hindlimbs, produced by transplanting hindlimb buds of chick embryos in place of forelimb buds, was quantified by analyzing the number and temporal distribution of spontaneous limb movements. Brachially innervated hindlimbs exhibited normal motility until E10 but thereafter became significantly less active than normal limbs and the limb movements were more randomly distributed. Contrary to the findings with axolotls and frogs, functional interaction between brachial motoneurons and hindlimb muscles cannot be sustained in the chick embryo. Dysfunction is first detectable at E10 and progresses to near total immobility by E20 and is associated with joint ankylosis and muscular atrophy. Although brachially innervated hindlimbs were virtually immobile by the time of hatching (E21), they produced strong movements in response to electrical stimulation of their spinal nerves, suggesting a central rather than peripheral defect in the motor system. The extent of motoneuron death in the brachial spinal cord was not significantly altered by the substitution of the forelimb bud with the hindlimb bud, but the timing of motoneuron loss was appropriate for the lumbar rather than brachial spinal cord, indicating that the rate of motoneuron death was dictated by the limb. Measurements of nuclear area indicated that motoneuron size was normal during the motoneuron death period (E6-E10) but the nuclei of motoneurons innervating grafted hindlimbs subsequently became significantly larger than those of normal brachial motoneurons. Although the muscle mass of the grafted hindlimb at E18 was significantly less than that of the normal hindlimb (and similar to that of a normal forelimb), electronmicroscopic examination of the grafted hindlimbs and brachial spinal cords of E20 embryos revealed normal myofiber and neuromuscular junction ultrastructure and a small increase in the number of axosomatic synapses on cross-sections of motoneurons innervating grafted hindlimbs compared to motoneurons innervating normal forelimbs. The anatomical data indicate that, rather than being associated with degenerative changes, the motor system of the brachial hindlimb of late-stage embryos is intact, but inactive. © 1993 John Wiley & Sons, Inc.     

Human body proportions explained on the basis of biomechanical principles

On the basis of theoretical biomechanics and of experiments, we investigated the mechanical requirements to which the body of a bipedally walking primate is subject, and the possibilities to meet these requirements with a minimum amount of energy. The least energy-consuming adaptation is clearly a body shape favourable for the preferred locomotion. Some characteristics of human body shape, in particular its proportions, could be identified as advantageous for fulfilling obvious biological roles or mechanical necessities. The characteristic length and the extended position of human hindlimbs make walking faster without additional input of energy. Mass distribution on the hindlimbs reduces the energy necessary for accelerating the swing limb after liftoff and for decelerating the swing limb before the heelstrike. Length and mass distribution in the forelimb gives it a pendulum length comparable to that of the hindlimb, so that both extremities swing at the same frequency. This swinging of the forelimbs counters in part the movements exerted by the moved hindlimbs on the trunk. The elongate and slim shape of the trunk provides great mass moments of inertia and that means stability against being flexed ventrally and dorsally by the forward and rearward movements of the heavy and long hindlimbs. Shoulder breadth in combination with the shallow shape of the thorax yield higher mass moments of inertia against the rotation of the trunk about a vertical axis than a cylindrical trunk shape. Further elongation of the hindlimbs is limited by the energy necessary for acceleration and deceleration, as well as for lifting them during the swing phase. In addition, the reaction forces exerted by the hindlimbs would expose the trunk to undue excursions if the proportions trunk length/limb length or trunk mass/limb mass would decrease. The above-noted kinetic requirements are partly in line, partly in conflict with the requirements of statics.     

SPEED AND STAMINA TRADE-OFF IN LACERTID LIZARDS

Abstract.— Morphological and physiological considerations suggest that sprinting ability and endurance capacity put conflicting demands on the design of an animal's locomotor apparatus and therefore cannot be maximized simultaneously. To test this hypothesis, we correlated size‐corrected maximal sprint speed and stamina of 12 species of lacertid lizards. Phylogenetically independent contrasts of sprint speed and stamina showed a significant negative relationship, giving support to the idea of an evolutionary trade‐off between the two performance measures. To test the hypothesis that the trade‐off is mediated by a conflict in morphological requirements, we correlated both performance traits with snout‐vent length, size‐corrected estimates of body mass and limb length, and relative hindlimb length (the residuals of the relationship between hind‐ and forelimb length). Fast‐running species had hindlimbs that were long compared to their forelimbs. None of the other size or shape variables showed a significant relationship with speed or endurance. We conclude that the evolution of sprint capacity may be constrained by the need for endurance capacity and vice versa, but the design conflict underlying this trade‐off has yet to be identified.     

How the cubic measure of the jumping style of the prosimian and its proportions is determined

How does body size determine the locomotor performance and proportions of leapers? In an analysis of the mechanics of leaping we derived two principles that explain the kinematic and morphological differences between leaping prosimian primates of different body size. 1. In small animals, the distance through which the body can be accelerated during take-off, and the time available for acceleration, are short. In small-bodied leapers we therefore find adaptations that increase the distance or length of time for propulsion and maximize speed. These are: great angular excursions at the joints of the hindlimb, long load arms of body weight and short power arms for the muscles, elongated hindlimbs with a disproportionate lengthening of the distal segments, and additional joints in the tarsus. 2. With increasing body size, the time for accelerating the body is no longer a problem. Instead, the ratio of muscle force available for acceleration to mass to be accelerated is unfavorable. Accordingly, large-bodied leapers have adaptations that allow optimal use of available muscle force. These include: acceleration in energetically profitable joint positions, avoidance of acute joint angles especially at the distal joints (where the muscles work against the highest percentage of body mass), only moderate elongation of the hindlimbs with rather short distal segments, and long lever arms of those muscles that extend the hindlimb joints. In addition, take-offs of the larger-bodied leapers are characterized by a regularly occurring arm swing movement, thus making additional use of nonhindlimb muscles for acceleration. The mass-dependent differences in forces and velocities have consequences for the energy budget. As the muscles of the small species must contract very rapidly against high loads, they consume more energy per unit of mechanical work. It is not possible to optimize speed and force in the same animal. Body size in conjunction with the laws of mechanics determines how maximum leaping potential will be realized.     

IMU: Inertial sensing of vertical CoM movement

The purpose of this study was to use a quaternion rotation matrix in combination with an integration approach to transform translatory accelerations of the centre of mass (CoM) from an inertial measurement unit (IMU) during walking, from the object system onto the global frame. Second, this paper utilises double integration to determine the relative change in position of the CoM from the vertical acceleration data. Five participants were tested in which an IMU, consisting of accelerometers, gyroscopes and magnetometers was attached on the lower spine estimated centre of mass. Participants were asked to walk three times through a calibrated volume at their self-selected walking speed. Synchronized data were collected by an IMU and an optical motion capture system (OMCS); both measured at 100 Hz. Accelerations of the IMU were transposed onto the global frame using a quaternion rotation matrix. Translatory acceleration, speed and relative change in position from the IMU were compared with the derived data from the OMCS. Peak acceleration in vertical axis showed no significant difference (p?0.05). Difference between peak and trough speed showed significant difference (p<0.05) but relative peak-trough position between the IMU and OMCS did not show any significant difference (p?0.05). These results indicate that quaternions, in combination with Simpsons rule integration, can be used in transforming translatory acceleration from the object frame to the global frame and therefore obtain relative change in position, thus offering a solution for using accelerometers in accurate global frame kinematic gait analyses.     

Mechanics of torque generation during quadrupedal arboreal locomotion

Quadrupedal animals moving on arboreal substrates face unique challenges to maintain stability. The torque generated by the limbs around the long axis of a branch during locomotion may clarify how the animals remain stable on arboreal supports. We sought to determine what strategy gray short-tailed opossums (Monodelphis domestica) use to exert torque and avoid toppling. The opossums moved across a branch trackway about half the diameter of their bodies. Part of the trackway was instrumented to measure substrate reaction forces and torque around the long axis of the branch. Kinematic analysis was used to estimate the center of pressure of the manus and pes; from center of pressure and vertical and mediolateral forces, the torque generated by substrate reaction forces versus muscular effort could be determined. Forelimbs generated significantly greater torque than hindlimbs, which is probably explained by the greater weight-bearing role of the forelimbs. Fore- and hindlimbs generated torque in opposite directions because contralateral fore- and hindlimbs typically contacted the branch. Torque generated by muscular effort, however, was often in the same direction in both fore- and hindlimbs. The muscle-generated torque is likely the result of mediolateral movement of the center of mass caused by mediolateral undulation of the torso. These results bear an important implication for the study of arboreal locomotion: center of mass dynamics are at least as important as static positions. M. domestica is a good representative for a primitive mammal, and comparisons with arboreal specialists will shed light on how proficient arboreal locomotion evolved.     

Effect of hindlimb unloading on interlimb coordination during treadmill locomotion in the rat

Effects of hindlimb unloading on interlimb coordination were examined in adult rats walking on a treadmill at moderate speed. In the first group of animals, the electromyographic activity (EMG) of soleus muscle of both hindlimbs was recorded after 7 and 14 days of unloading. In the second group, the EMG was recorded daily until the 14th day of unloading. The general organization of locomotion was preserved in the two groups whatever the duration of the unloading. The step cycles of the two hindlimbs were always strictly alternating. However, the locomotor pattern was very irregular. A lateral instability was observed. It was accompanied by an abduction of the hindlimbs, and frequent hyperextensions of the ankle when walking. The EMG analysis showed an increase in step cycle duration and in coactivation duration of the soleus muscles (i.e. in the double stance duration). In the rats recorded daily, mean EMG was dramatically reduced the 1st day of unloading, suggesting a decrease in the neural drive. Taken together, these data indicate that 14 days of hindlimb unloading can alter the neuromuscular pattern during locomotion. It is proposed that these changes are related to changes in the peripheral sensory information. Accepted: 29 June 1998     

Development of an advanced mechanised gait trainer, controlling movement of the centre of mass, for restoring gait in non-ambulant subjects.

The study aimed at further development of a mechanised gait trainer which would allow non-ambulant people to practice a gait-like motion repeatedly. To simulate normal gait, discrete stance and swing phases, lasting 60% and 40% of the gait cycle respectively, and the control of the movement of the centre of mass were required. A complex gear system provided the gait-like movement of two foot plates with a ratio of 60% to 40% between the stance and swing phases. A controlled propulsion system adjusted its output according to patient's efforts. Two eccenters on the central gear controlled phase-adjusted the vertical and horizontal position of the centre of mass. The patterns of sagittal lower limb joint kinematics and of muscle activation of a normal subject were similar when using the mechanised trainer and when walking on a treadmill. A non-ambulatory hemiparetic subject required little help from one therapist on the gait trainer, while two therapists supported treadmill walking. Gait movements on the trainer were highly symmetrical, impact-free, and less spastic. The weight-bearing muscles were activated in a similar fashion during both conditions. The vertical displacement of the centre of mass was bi-instead of mono-phasic during each gait cycle on the new device. In conclusion, the gait trainer allowed wheelchair-bound subjects the repetitive practice of a gait-like movement without overstraining therapists.     

Radiation inactivation studies on Photosystem I. Functional sizes of electron-transport reactions

Radiation inactivation studies on the functional size of electron-transport processes in the Photosystem I reaction-centre complex showed the following characteristics. (1) The molecular mass required for electron transport from P-700 to iron-sulphur centre A was below 40 kDa. (2) Independent inactivation of iron-sulphur centres A and B was observed indicating their location on separate polypeptides. (3) The molecular mass of the polypeptides containing iron-sulphur centres A and B were 5–10 kDa based on a linear electron-transfer chain or 15–20 and 5–10 kDa (centre B) based on a branched chain. (4) A reaction centre ‘core’ containing the electron carriers for electron transport from P-700 to iron-sulphur centre X was indicated. These observations are discussed in comparison to current ideas on the polypeptide composition of the Photosystem I reaction centre. It is concluded that the radiation inactivation technique did not measure the size of Photosystem I polypeptides binding chlorophyll accounting for the small overall target size. The observed functional size came mostly from inactivation of the iron-sulphur centres showing that they are located on separate polypeptides.     

Bone density, breaking force and leg muscle mass as functions of weight in bipedal rats

Femur density, femur breaking force and muscle weight on the hind limbs of normal and bipedal rats have been measured. The bipeds had more muscle on the hindlimbs than controls. Increasing muscle mass was associated with increasing femur density and breaking force. It is concluded that weight bearing influences bone density and breaking force through muscle mass.     

An objective classification system of air mass types for Szeged, Hungary, with special attention to plant pollen levels

This paper discusses the characteristic air mass types over the Carpathian Basin in relation to plant pollen levels over annual pollination periods. Based on the European Centre for Medium-Range Weather Forecasts dataset, daily sea-level pressure fields analysed at 00 UTC were prepared for each air mass type (cluster) in order to relate sea-level pressure patterns to pollen levels in Szeged, Hungary. The database comprises daily values of 12 meteorological parameters and daily pollen concentrations of 24 species for their pollination periods from 1997 to 2001. Characteristic air mass types were objectively defined via factor analysis and cluster analysis. According to the results, nine air mass types (clusters) were detected for pollination periods of the year corresponding to pollen levels that appear with higher concentration when irradiance is moderate while wind speed is moderate or high. This is the case when an anticyclone prevails in the region west of the Carpathian Basin and when Hungary is under the influence of zonal currents (wind speed is high). The sea level pressure systems associated with low pollen concentrations are mostly similar to those connected to higher pollen concentrations, and arise when wind speed is low or moderate. Low pollen levels occur when an anticyclone prevails in the region west of the Carpathian Basin, as well as when an anticyclone covers the region with Hungary at its centre. Hence, anticyclonic or anticyclonic ridge weather situations seem to be relevant in classifying pollen levels.