Evolutionary Biomechanics

The Carnivora spans the largest ecological and body size diversity of any mammalian order, making it an ideal basis for studies of evolutionary ecology and functional morphology. For animals with different feeding ecologies, it may be expected that bite force represents an important evolutionary adaptation, but studies have been constrained by a lack of bite force data. In this study we present predictions of bite forces for 151 species of extant carnivores, comprising representatives from all eight families and the entire size and ecological spectrum within the order. We show that, when normalized for body size, bite forces differ significantly between the various feeding categories. At opposing extremes and independent of genealogy, consumers of tough fibrous plant material and carnivores preying on large prey both have high bite forces for their size, while bite force adjusted for body mass is low among specialized insectivores. Omnivores and carnivores preying on small prey have more moderate bite forces for their size. These findings indicate that differences in bite force represent important adaptations to and indicators of differing feeding ecologies throughout carnivoran evolution. Our results suggest that the incorporation of bite force data may assist in the construction of more robust evolutionary and palaeontological analyses of feeding ecology.

Tetrapods possess up to five morphologically distinct vertebral series: cervical, thoracic, lumbar, sacral and caudal. The evolution of axial regionalization has been linked to derived Hox expression patterns during development and the demands of weight-bearing and walking on land. These evolutionary and functional explanations are supported by an absence of similar traits in fishes, living and extinct. Here, I show that, Tarrasius problematicus, a marine ray-finned fish from the Mississippian (Early Carboniferous; 359–318 Ma) of Scotland, is the first non-tetrapod known to possess tetrapod-like axial regionalization. Tarrasius exhibits five vertebral regions, including a seven-vertebrae ‘cervical’ series and a reinforced ‘sacrum’ over the pelvic area. Most vertebrae possess processes for intervertebral contact similar to tetrapod zygapophyses. The fully aquatic Tarrasius evolved these morphologies alongside other traits convergent with early tetrapods, including a naked trunk, and a single median continuous fin. Regional modifications in Tarrasius probably facilitated pelagic swimming, rather than a terrestrial lifestyle or walking gait, presenting an alternative scenario for the evolution of such traits in tetrapods. Axial regionalization in Tarrasius could indicate tetrapod-like Hox expression patterns, possibly representing the primitive state for jawed vertebrates. Alternately, it could signal a weaker relationship, or even a complete disconnect, between Hox expression domains and vertebrate axial plans.

Some tetrapods hang upside down from tree branches when moving horizontally. The ability to walk in quadrupedal suspension has been acquired independently in at least 14 mammalian lineages. During the stance (supportive) phase of quadrupedal suspension, the elbow joint flexor muscles (not the extensors as in upright vertebrates moving overground) are expected to contract to maintain the flexed limb posture. Therefore muscular control in inverted, suspended quadrupeds may require changes of muscle control, and even morphologies, to conditions opposite to those in upright animals. However, the relationships between musculoskeletal morphologies and elbow joint postures during the stance phase in suspended quadrupeds have not been investigated. Our analysis comparing postures and skeletal morphologies in Choloepus (Pilosa), Pteropus (Chiroptera), Nycticebus (Primates) and Cynocephalus (Dermoptera) revealed that the elbow joints of these animals were kept at flexed angles of 70–100 ° during the stance phase of quadrupedal suspension. At these joint angles the moment arms of the elbow joint flexors were roughly maximized, optimizing that component of antigravity support. Our additional measurements from various mammalian species show that suspended quadrupeds have relatively small extensor/flexor ratios in both muscle masses and maximum moment arms. Thus, in contrast to the pattern in normal terrestrial quadrupeds, suspended quadrupeds emphasize flexor over extensor muscles for body support. This condition has evolved independently multiple times, attendant with a loss or reduction of the ability to move in normal upright postures.

Jill E. Scott, John D. Polk. Dept. of Anthropology, University of Illinois at Urbana-Champaign.

The purpose of this project is to provide a preliminary experimental test of a biomechanical model that relates the shock-absorbing characteristics of intervertebral disks to their cross-sectional area.  This has implications for studying the evolution of bipedal distance running in hominins since vertebral body size, and hence, intervertebral disk size increases with the origin of Homo erectus, the taxon believed to be the first endurance runner; and because impact force magnitudes differ dramatically between walking and running.  An in vitro study was conducted using six simulated intervertebral disks of varying area, but otherwise identical material properties.  Each disk was subjected to impact loads and transmitted force characteristics were measured using a force platform (AMTI Inc. Watertown, MA).  Specific measurements include the peak force transmitted through each disk, the duration of time that the load remained in contact with each disk, and the amount of energy absorbed by each disk.  The results show that, for a constant impact load, the energy absorbed by each disk did not increase significantly with increasing disk area.  Impact forces transmitted through the disk increased with disk area, while the duration of the impact load decreased with increasing disk area.  These results suggest that increasing disk area alone will not confer an advantage for impact force absorption.  Our model suggests that concomitant changes in disk material properties and thickness may be more effective at moderating the transmission of impact forces, and these changes should be expected in the vertebral columns of bipeds adapted to distance running.

Intensive artificial selection has led to the production of the modern broiler chicken, which over the last few decades has undergone a dramatic increase in growth rate and noticeable changes in body conformation. Unfortunately, this has been associated with musculoskeletal abnormalities which have altered the walking ability of these birds, raising obvious welfare concerns, as well as causing economic losses. Here we present a comparative study of ancestral and derived muscle anatomy in chickens to begin to tease apart how evolutionary alterations of muscle form in chickens have influenced their locomotor function and perhaps contributed to lameness. We measured the muscle architectural properties of the right pelvic limb in 50 birds, including the Giant Junglefowl, a commercial strain broiler and four pureline commercial broiler breeder lines (from which the broiler populations are derived) to identify which features of the broiler’s architectural design have diverged the most from the ancestral condition. We report a decline in pelvic limb muscle mass in the commercial line birds that may compromise their locomotor abilities because they carry a larger body mass. This greater demand on the pelvic limb muscles has mostly led to changes in support at the hip joint, revealing significantly larger abductors and additionally much larger medial rotators in the broiler population. Differences were seen within the commercial line bird populations, which are likely attributed to different selection pressures and may reflect differences in the walking ability of these birds. In addition, Junglefowl seem to have both greater force-generating capabilities and longer, presumably faster contracting muscles, indicative of superior musculoskeletal/locomotor function. We have provided baseline data for generating hypotheses to investigate in greater depth the specific biomechanical constraints that compromise the modern broiler’s walking ability and propose that these factors should be considered in the selection for musculoskeletal health in the chickens of the future. Our new anatomical data for a wide range of domestic and wild-type chickens is useful in a comparative context and for deeper functional analysis including computer modelling/simulation of limb mechanics.

Intensive artificial selection has led to the production of the modern broiler chicken, which over the last few decades has undergone a dramatic increase in growth rate and noticeable changes in body conformation. Unfortunately, this has been associated with musculoskeletal abnormalities which have altered the walking ability of these birds, raising obvious welfare concerns, as well as causing economic losses. Here we present a comparative study of ancestral and derived muscle anatomy in chickens to begin to tease apart how evolutionary alterations of muscle form in chickens have influenced their locomotor function and perhaps contributed to lameness. We measured the muscle architectural properties of the right pelvic limb in 50 birds, including the Giant Junglefowl, a commercial strain broiler and four pureline commercial broiler breeder lines (from which the broiler populations are derived) to identify which features of the broiler’s architectural design have diverged the most from the ancestral condition. We report a decline in pelvic limb muscle mass in the commercial line birds that may compromise their locomotor abilities because they carry a larger body mass. This greater demand on the pelvic limb muscles has mostly led to changes in support at the hip joint, revealing significantly larger abductors and additionally much larger medial rotators in the broiler population. Differences were seen within the commercial line bird populations, which are likely attributed to different selection pressures and may reflect differences in the walking ability of these birds. In addition, Junglefowl seem to have both greater force-generating capabilities and longer, presumably faster contracting muscles, indicative of superior musculoskeletal/locomotor function. We have provided baseline data for generating hypotheses to investigate in greater depth the specific biomechanical constraints that compromise the modern broiler’s walking ability and propose that these factors should be considered in the selection for musculoskeletal health in the chickens of the future. Our new anatomical data for a wide range of domestic and wild-type chickens is useful in a comparative context and for deeper functional analysis including computer modelling/simulation of limb mechanics.

The problem that modern philosophy of science has with teleological explanations of natural objects is that they are by no means epistemologically justified, and still we tend to explain natural objects by referring to their functions as if they were causally determined by their ends. This paper's purpose is to show how Darwin's theory of natural selection made possible to consider organisms' adaptation to their environment without reference to an intelligent designer.

We developed a method based on interactive B-spline solids for estimating and visualizing biomechanically important parameters for animal body segments. Although the method is most useful for assessing the importance of unknowns in extinct animals, such as body contours, muscle bulk, or inertial parameters, it is also useful for non-invasive measurement of segmental dimensions in extant animals. Points measured directly from bodies or skeletons are digitized and visualized on a computer, and then a B-spline solid is fitted to enclose these points, allowing quantification of segment dimensions. The method is computationally fast enough so that software implementations can interactively deform the shape of body segments (by warping the solid) or adjust the shape quantitatively (e.g., expanding the solid boundary by some percentage or a specific distance beyond measured skeletal coordinates). As the shape changes, the resulting changes in segment mass, center of mass (CM), and moments ...

Many bones are supported internally by a latticework of trabeculae. Scaling of whole bone length and diameter has been extensively investigated, but scaling of the trabecular network is not well characterized. We analysed trabecular geometry in the femora of 90 terrestrial mammalian and avian species with body masses ranging from 3 g to 3400 kg. We found that bone volume fraction does not scale substantially with animal size, while trabeculae in larger animals' femora are thicker, further apart and fewer per unit volume than in smaller animals. Finite element modelling indicates that trabecular scaling does not alter the bulk stiffness of trabecular bone, but does alter strain within trabeculae under equal applied loads. Allometry of bone's trabecular tissue may contribute to the skeleton's ability to withstand load, without incurring the physiological or mechanical costs of increasing bone mass.