Economy and Endurance in Human Evolution.

The evolutionary pressures shaping humans' unique bipedal locomotion have been a focus of research since Darwin, but the origins of humans' economical walking gait and endurance running capabilities remain unclear. Here, I review the anatomical and physiological determinants of locomotor economy (e.g., limb length and posture) and endurance (e.g., muscle volume and fiber type) and investigate their development in the hominin fossil record. The earliest hominins were bipedal but retained ape-like features in the hind limb that would have limited their walking economy compared to living humans. Moreover, the evolution of bipedalism and the loss of the forelimbs in weight support and propulsion would have reduced locomotor endurance in the earliest hominins and likely restricted ranging. Australopithecus evinced longer hind limbs, extended limb posture, and a stiff midfoot, suggesting improved, human-like economy, but were likely still limited in their endurance compared to modern humans. The appearance of skeletal traits related to endurance (e.g., larger limb joints, spring-like plantar arch) in Homo was somewhat mosaic, with the full endurance suite apparent only ∼1 million years ago. The development of endurance capabilities in Homo appears to parallel the evolutionary increase in brain size, cognitive sophistication, and metabolic rate.

An interspecific assessment of Bergmann's rule in 22 mammalian families.

BACKGROUND: Bergmann's rule proposes that animals in cold habitats will be larger than those in warm habitats. This prediction has been tested thoroughly at the intraspecific level, but few studies have investigated the hypothesis with interspecific data using phylogenetic comparative approaches. Many clades of mammals have representatives in numerous distinct biomes, making this order highly suitable for a large-scale interspecific assessment of Bergmann's rule. Here, we evaluate Bergmann's rule within 22 mammalian families-with a dataset that include ~35 % of all described species-using a phylogenetic comparative approach. The method is based on an Ornstein-Uhlenbeck model of evolution that allows for joint estimation of adaptation and constraints (phylogenetic inertia) in the evolution of a trait. We use this comparative method to investigate whether body mass evolves towards phenotypic optima that are functions of median latitude, maximum latitude or temperature. We also assess the closely related Allen's rule in five families, by testing if relative forelimb length evolves as a function of temperature or latitude. RESULTS: Among 22 mammalian families, there was weak support for Bergmann's rule in one family: A decrease in temperature predicted increased body mass in Canidae (canids). We also found latitude and temperature to significantly predict body mass in Geomyidae (pocket gophers); however, the association went in the opposite direction of Bergmann's predictions. Allen's rule was supported in one of the five examined families (Pteropodidae; megabats), but only when forelimb length evolves towards an optimum that is a function of maximum latitude, not median latitude or temperature. CONCLUSIONS: Based on this exhaustive assessment of Bergmann's rule, we conclude that factors other than latitude and temperature are the major drivers of body mass evolution at the family level in mammals.

Ex Vivo Assessment of an Ultrasound-Guided Injection Technique of the Navicular Bursa in the Horse.

Synovitis of the navicular bursa is common in performance horses. The objective of this study was to describe an ultrasound-guided technique to inject a distended navicular bursa and to evaluate its feasibility for use by a clinician not trained in the technique. Twenty distal limbs of horses of various breeds and sizes were used. To produce synovial distension, the navicular bursa of each limb was injected with contrast medium using a lateral approach and radiography was performed to confirm that the contrast medium was distending the bursa. The digit was positioned with the distal interphalangeal joint in hyperextension. A microconvex ultrasound probe was placed in the hollow of the pastern, palmar to the middle phalanx and the region was assessed in a transverse plane slightly oblique to the horizontal plane. The ultrasound probe was rotated to visualize both the lateral and medial recesses and to select which side was more distended to inject. A 21G 0.8 × 50 mm needle was inserted abaxially to the probe in the plane of the ultrasound beam into the proximal recess of this navicular bursa and a methylene blue solution was injected. Following injection, dissection was performed to assess whether the navicular bursa had been successfully injected. This ultrasound-guided technique was reliably performed with a success rate of 68%. The success of injection is influenced by hyperextension of the foot, quality of ultrasound images and degree of distension of the bursa.

CT assessment of the influence of dynamic loading on physiological incongruency of the canine elbow.

OBJECTIVES: To present a novel technique for loading of the canine elbow joint and to quantify changes in congruency with increasing load using computed tomography. MATERIALS AND METHODS: Five pairs of thoracic limbs were mounted at a mid stance angle in a custom made jig. Elbow joints were loaded to 0, 33, 66 and 100% of total individual cadaver bodyweight. At each load computed tomography of the elbow was performed. Joint space measurement was performed on sagittal plane central, lateral and medial compartment images at humero-radial (R1 , R2 , R3 ), humero-ulnar (U1 , U2 ) and radio-ulnar loci. The effect of loading on joint spaces was assessed (P<0.05). RESULTS: With increasing load; for central an increase in R1 and radio-ulnar distance and decreased R3 occurred; for medial R1 increased; and for lateral R1 and radio-ulnar distance increased. The largest increases were seen in the lateral compartment. CLINICAL SIGNIFICANCE: Significant changes in humero-radio-ulnar congruency occurred suggesting pronation of the radius with respect to the ulna was induced during loading. This movement may influence the load experienced by the medial coronoid process and could play a role in the aetiopathogenesis of medial coronoid process disease.

Inter- and intratester reliability of anthropometric assessment of limb circumference in labrador retrievers.

OBJECTIVE: To report a standardized method of, and determine inter- and intratester reliability for, anthropometric assessment of limb circumference in dogs. STUDY DESIGN: Prospective blinded study. ANIMALS: Labrador Retrievers (n = 20). METHOD: Unsedated dogs were manually restrained in lateral recumbency and triplicate measurements of limb circumference at the level of the proximal antebrachium, mid brachium, proximal crus, and mid-thigh were made using the Gulick II tape measure in the morning and afternoon of the same day. Observers were blinded to measurements made during each occasion and those made by co-observers. Estimates of inter- and intratester reliability were made for first and mean measurements using intra-class correlation coefficients (ICC). RESULTS: Measurements of the proximal antebrachium were made with moderate to fair intratester reliability by all observers with ICC's ranging from 0.68-0.78 (1st measurement) and 0.67-0.78 (mean measurement), and moderate to fair intertester reliability with ICC's of 0.66-0.68 (1st measurement) and 0.70-0.72 (mean measurement). Measurements of the brachium, crus, and thigh typically had poor inter- and intratester reliability, ICC < 0.5. CONCLUSION: Using the described method of muscle measurement in Labrador Retrievers only measurement of the proximal antebrachium was reliable; a single (1st) measurement was as reliable as using the mean of triplicate measurements.

Assessment of ultrasonographic morphometric measurements of digital flexor tendons and ligaments of the palmar metacarpal region in Icelandic Horses.

OBJECTIVE: To obtain morphometric values for the superficial digital flexor tendon, deep digital flexor tendon, accessory ligament of the deep digital flexor muscle, and suspensory ligament in the palmar metacarpal region of Icelandic Horses. ANIMALS: 50 nonlame Icelandic Horses in training. PROCEDURES: Horses included 2 stallions, 30 geldings, and 18 mares from 4 to 20 years of age with a body mass index from 149.1 to 250.11 kg/m2. Transverse ultrasonographic images were obtained with an 8- to 10-MHz linear transducer and a standoff pad. In both forelimbs, the cross-sectional area, circumference, dorsopalmar width, and lateromedial width were measured 3 times at 5 regions of interest (ROIs). RESULTS: The coefficient of variation for all measurements of each ROI was <5%. Comparisons were performed among and within structures and for each variable at all ROIs. Comparisons among horses revealed homogeneity because no significant influences of age, sex, height at the withers, or body mass index were found. Additionally, a characteristic skin condition interfering with ultrasonographic examination was observed. CONCLUSIONS AND CLINICAL RELEVANCE: The morphometric values of the structures examined were consistent with those reported for other breeds, although some differences were observed.

Assessment of the geometry of the cubital (elbow) joint of dogs by use of magnetic resonance imaging.

OBJECTIVE: To assess joint geometry and highlight potential evidence of physiologic incongruity in cubital (elbow) joints of large, small, and chondrodystrophic breeds of dogs. SAMPLE POPULATION: Nonarthritic elbow joints obtained from cadavers of mature dogs of large breeds (n = 19), small breeds (14), or chondrodystrophic breeds (8). PROCEDURE: Magnetic resonance imaging was used to quantify interosseous gaps at 6 defined positions and 2 sagittal planes of the humeroantebrachial region by use of a fat-suppressed 3-dimensional gradient-echo sequence. RESULTS: Interosseous gaps in the more medially located sagittal plane were significantly narrower at the level of the anconeal process than at any other position of the joint in large-breed dogs, compared with results for small- or chondrodystrophic-breed dogs. In both sagittal planes, the gaps were significantly wider at the center of the ulnar trochlear notch than at any other position. Significant correlation between body weight and width of the interosseous gaps was found only in large-breed dogs and was found in both sagittal planes at the center of the ulnar trochlear notch (r, 0.834 and r, 0.680, respectively). CONCLUSIONS AND CLINICAL RELEVANCE: In large-breed dogs, the interosseous gap was especially narrow at the level of the anconeal process and its diametric position. This suggests less ability to compensate intra-articular steps in dogs with short ulna syndrome, which predisposes to failure of the union between the anconeal process and olecranon. Geometric incongruity may be regarded as clinically normal in nonarthritic humeroulnar joints. The degree of geometric incongruity increases with body weight.

Use of reconstructed computed tomography for the assessment of joint spaces in the canine elbow.

OBJECTIVES: To assess the accuracy of reconstructed computed tomography for imaging canine elbow joint spaces. METHODS: Computed tomography scans of eight cadaveric elbows were obtained and reconstructed computed tomography images were formatted in the dorsal and sagittal planes. Humeroradial and humeroulnar joint space measurements were obtained from these images. Intra-observer and inter-observer variations in joint space measurements were assessed, as was the effect of specimen positioning (inter-image variation). After freezing, four elbows were sectioned in the dorsal plane and four in the sagittal plane. In addition to visual comparison of the frozen sections with reconstructed computed tomography images, joint space measurements were obtained from frozen sections and compared with those from reconstructed computed tomography images. Variation was assessed using statistical calculations and graphical techniques. RESULTS: Both inter-image and intra-observer analyses revealed good agreement and low variation between data sets. Inter-observer correlation was only moderate, though variation was low. Visually, the reconstructed computed tomography images accurately reflected the frozen section anatomy. Agreement between frozen section and reconstructed computed tomography joint space measurements was good, with minimal variation. CLINICAL SIGNIFICANCE: These results indicate that reconstructed computed tomography is capable of accurately imaging elbow joint spaces and precise joint space measurements can be obtained. Reconstructed computed tomography may be useful for determining joint space measurements and detecting elbow incongruencies in dogs with elbow dysplasia.

The cost of transport in an extended trot.

We hypothesised that trotters during an extended trot have lower energetic costs of locomotion (CT) than horses not bred for this behaviour. VO2 was measured as a function of speed in 7 Arabian horses (3 trained to extend their trotting speeds) and in 2 horses, of similar mass, bred to trot (Hackney). Both oxygen consumption and CT increased with speed and there was, contrary to our hypothesis, no difference between breeds. In Arabians at 6.5 m/s, CT had increased 25% above the CT at 5.0 m/s (normal transition speed). For Hackneys at 6.8 m/s, the CT was almost 35% higher. Stride frequencies increased linearly in all horses up to 5.0 m/s. At the canter at 5.0 m/s, the frequency increased 9% to 111 strides/min, but then increased minimally with speed. In the Hackneys and the Arabians that extended the trot, stride frequencies were approximately 102 and did not increase with speed. Stride length (SL) increased linearly with speed in both trotting and cantering horses, and cantering SL were lower than trotting (at 5.0 m/s, SL for trotting = 3.04 m and for cantering SL = 2.68 m). There were no differences between breeds in stride frequency or stride length. Extending the trot can have profound energetic requirements that could limit athletic performance and may lead to increased concussive impact on the limbs.

Gait, estimated net cost of transport and heat production at different speeds in three-day-event horses.

Heart rate and gait characteristics (stride length and frequency) were studied in 6 horses subjected to a standardised incremental exercise test, involving moving at the trot and increasing speeds up to a fast gallop and subsequently during the steeplechase phase of a 3-day-event. The studies were performed in hot conditions. Appropriate scaling, based on hindleg length (hh), stride length (L), stride frequency (f) and speed (Sp) for nondimensional stride length (lambda = (L/hh), nondimensional stride frequency (phi = f(hh/g)1/2) and nondimensional velocity (û = Sp/(ghh)1/2), where g is the gravitational acceleration, demonstrated that there were no major differences in characteristics over the full range studied lambda = 2.3 û0.68. However, there were subtle differences in some horses that could endow a benefit in locomotory efficiency when compared to others exercising at the same absolute speed. There were clear changes in the relationship between nondimensional stride length and frequency with increasing speed, from trot to canter (at û approximately 1.4) and to full gallop (at û approximately 2.3); when trotting, lambda was less than 2.2 and the transition from canter to gallop took place at lambda approximately 3.2. The cost of transport/kg/m, estimated from the heart rates measured continuously during each study, decreased with increasing speed and bodyweight. In some animals, there appeared to be a weak minimum around the canter-gallop transition speed. When interpreted as oxygen cost, using published values for the oxygen consumption-heart rate relationship, the cost fell from an average of 0.201 ml/kg/m at the trot to 0.161 mlO2/kg/m when galloping during the incremental exercise tests. During the steeplechase, the cost was approximately 7.5% higher than at the same speed in the exercise tests; this was probably due to jumping effort. Estimated power consumption increased linearly with speed. In the steeplechase, power consumption was also 7.5% higher than during the exercise tests at the same absolute speed; this was equivalent to an average rate of heat production of 346 kcal/min (24 kW) or 59.5 kcal/min/m2 of the measured body surface area.