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David R Carrier

Publications and source records attributed to David R Carrier.

14 recordsLinked to original sources
PubMed2007

Ground forces applied by galloping dogs.

The gallop differs from most other quadrupedal gaits in that each limb plays a unique role. This study compares the ground forces applied by the four limbs and uses force differences between limbs to address the question of why the gallop is the fastest quadrupedal gait. Individual ground forces were recorded from each of the four limbs as six dogs galloped down a runway at constant speed. Trials were videotaped at high speed using a camera positioned perpendicular to the runway, and velocity was measured using photosensors. The trailing forelimb applied greater peak vertical forces than the lead forelimb, however the vertical impulses from the two forelimbs were similar because the lead forelimb had a longer contact interval. The trailing forelimb and lead hindlimb applied greater peak accelerating forces and accelerating force impulses than their contralateral limbs despite their tendency to have shorter contact intervals. The accelerating impulse of both forelimbs combined did not differ significantly from that of both hindlimbs. The forelimbs applied a greater decelerating impulse than the hindlimbs, such that their net fore-aft impulse was decelerating whereas that of the hindlimbs was accelerating. The greater accelerating impulse applied by the trailing forelimb and greater decelerating impulse applied by the lead forelimb are consistent with the forelimbs acting as elastic struts rather than being actively retracted. In contrast, greater accelerating forces were produced by the lead hindlimb while the center of mass was lifted, suggesting that the hindlimbs are more actively extended or retracted during stance. The differences in ground forces measured between paired limbs suggest that the lead forelimb and trailing hindlimb are limited in their ability to apply forces by their positions in the stride cycle rather than by their muscular capacity. Although a bound or half-bound would allow more limbs to produce their maximal forces, a gallop may generate higher speeds because it is more efficient. Galloping could be more efficient than other gaits involving sagittal bending if the increased number of ground contact intervals decreased either the decelerating forces applied at the onset of ground contact or the vertical motion of the center of mass.

Animals↗
PubMed2006

The coupled evolution of breathing and locomotion as a game of leapfrog.

Because the increase in metabolic rate related to locomotor activity places demands on the cardiorespiratory apparatus, it is not surprising that the evolution of breathing and of locomotion are coupled. As the respiratory faculty becomes more refined, increasingly aerobic life strategies can be explored, and this activity is in turn expedited by a higher-performance respiratory apparatus. This apparent leapfrogging of respiratory and locomotor faculties begins in noncraniate chordates and continues in water-breathing and air-breathing vertebrates. Because both locomotor and cardiorespiratory activities are coordinated in the brain, neurological as well as biochemical coupling is evident. In spite of very different breathing mechanisms in various vertebrate groups, the basic respiratory control mechanisms appear to have been conserved, and respiratory-locomotor coupling is evident in all classes of vertebrates. Hypaxial body wall muscles that were strictly locomotor in fish have respiratory function in amniotes, but some locomotor function remains in all groups.

Animals↗
PubMed2006

Living history of physiology: Carl Gans.

In 2005, The American Physiological Society initiated The Living History of Physiology Project to recognize senior members who have made extraordinary contributions during their career to the advancement of the discipline and profession of physiology. Each Eminent Physiologist will be interviewed for archival purposes, and the video tape will be available from the American Physiological Society Headquarters. In addition, a biographical profile of the recipient will be published in Advances in Physiology Education.

History, 20th Century↗
PubMed2006

Locomotor function of the pectoral girdle 'muscular sling' in trotting dogs.

In therian mammals, gravitational and locomotor forces are transferred between the forelimb and trunk primarily, or entirely, through the muscles that connect the limb and trunk. Our understanding of this force transmission is based on analyses of shoulder anatomy and on a handful of descriptive electromyographic studies. To improve our understanding, we manipulated the locomotor forces of trotting dogs and monitored the resulting change in recruitment of five extrinsic muscles of the forelimb: m. serratus ventralis thoracis, m. serratus ventralis cervicis, m. pectoralis superficialis transversus, the anterior portion of the m. pectoralis profundus, and m. rhomboideus thoracis. Locomotor forces were modified as the dogs trotted at constant speed on a motorized treadmill by (1) adding mass to the trunk, (2) inclining the treadmill so that the dogs ran up and down hill, (3) adding mass to the wrists and (4) applying horizontally directed force to the trunk through a leash. These experiments indicate that the thoracic portion of the serratus ventralis muscle is the main antigravity muscle of the shoulder during trotting in dogs. Its activity increased when we added mass to the trunk and also when we ran the subjects downhill. In contrast, the cervical portion of the serratus ventralis did not show a consistent increase in activity in response to added mass. Instead, its activity increased when we ran the subjects up hill and added mass to their wrists, suggesting that it functions to stabilize the fulcrum of the forelimb in the cranial-caudal direction during active retraction of the forelimb. The thoracic portion of the rhomboideus muscle also appears to provide this cranial-caudal stabilization during active retraction of the forelimb. The force manipulations indicate that the transverse pectoralis muscle acts to both protract and retract the forelimb, depending on the position of the limb. In contrast, the anterior portion of the pectoralis profundus muscle acts as a retractor of the forelimb during the end of swing phase and the beginning of support phase. We found that adding mass to the trunk did not increase the activity of forelimb retractor muscles, suggesting that the ground reaction force vector passes through, or very near, the fulcrum of the shoulder during a trotting step. Whether or not the functions of these extrinsic appendicular muscles in dogs characterize therian mammals or represent specializations for high-speed, economical running remains to be determined.

Animals↗
PubMed2005

Interaction between the X chromosome and an autosome regulates size sexual dimorphism in Portuguese Water Dogs.

Size sexual dimorphism occurs in almost all mammals. In Portuguese Water Dogs, much of the difference in skeletal size between females and males is due to the interaction between a Quantitative Trait Locus (QTL) on the X-chromosome and a QTL linked to Insulin-like Growth Factor 1 (IGF-1) on the CFA 15 autosome. In females, the haplotype of CFA 15 resulting in small size is dominant. In males, the haplotype for large size is dominant. Females, homozygous at the CHM marker on the X chromosome and homozygous for the large size CFA 15 haplotype are, on average, as large as large males. However, all females that are heterozygous at the CHM marker are small, regardless of their CFA 15 genotype. This interaction suggests a genetic mechanism that in turn leads to a scenario for the evolution of size sexual dimorphism consistent with a proposal of Lande that sexual dimorphism can evolve because females secondarily become smaller than males as a consequence of natural selection for optimal size. Our results also can explain Rensch's Rule, which states that size is often positively correlated with the level of size sexual dimorphism.

Animals↗
PubMed2005

Genetics of canid skeletal variation: size and shape of the pelvis.

The mammalian skeleton presents an ideal system in which to study the genetic architecture of a set of related polygenic traits and the skeleton of the domestic dog (Canis familiaris) is arguably the best system in which to address the relationship between genes and anatomy. We have analyzed the genetic basis for skeletal variation in a population of >450 Portuguese Water Dogs. At this stage of this ongoing project, we have identified >40 putative quantitative trait loci (QTLs) for heritable skeletal phenotypes located on 22 different chromosomes, including the "X." A striking aspect of these is the regulation of suites of traits representing bones located in different parts of the skeleton but related by function. Here we illustrate this by describing genetic variation in postcranial morphology. Two suites of traits are involved. One regulates the size of the pelvis relative to dimensions of the limb bones. The other regulates the shape of the pelvis. Both are examples of trade-offs that may be prototypical of different breeds. For the size of the pelvis relative to limb bones, we describe four QTLs located on autosome CFA 12, 30, 31, and X. For pelvic shape we describe QTLs on autosome CFA 2, 3, 22, and 36. The relation of these polygenic systems to musculoskeletal function is discussed.

Animals↗
PubMed2004

Gear ratios at the limb joints of jumping dogs.

An increase in gear ratio of the limb extensor muscles during joint extension has been suggested to be a mechanism that facilitates optimal power production by skeletal muscles. The objectives of this study were to: (1) measure gear ratios at the wrist, elbow, shoulder, ankle, knee, and hip joints of jumping dogs, (2) compute the work performed by each of these joints, and (3) measure muscle shortening velocity for a joint exhibiting an increasing gear ratio during joint extension. The gear ratio out-lever was computed by dividing the ground reaction force (GRF) moment by the GRF, whereas the in-lever was directly measured as the perpendicular distance from the joint center to the line of action of the extensor muscle. In addition, changes in fascicle length were measured from the vastus lateralis muscle using sonomicrometry. Of the joints examined, only the gear ratios at the shoulder and knee joints increased during jumping in a manner that could facilitate peak power production of actively shortening muscles. The vastus lateralis was found to shorten at an average velocity of 3.20 muscle lengths per second. This is similar to estimates of shortening velocity that produce peak muscular power in mammals the size of dogs. Additionally, the knee extensors were found to produce a large proportion (26.6%) of the positive external work of the limbs. These observations suggest that dynamic gearing in jumping dogs may allow the extensor muscles of the knee joint to shorten in a way that maximizes their power production.

Animals↗
PubMed2004

Effects of mass distribution on the mechanics of level trotting in dogs.

The antero-posterior mass distribution of quadrupeds varies substantially amongst species, yet the functional implications of this design characteristic remain poorly understood. During trotting, the forelimb exerts a net braking force while the hindlimb exerts a net propulsive force. Steady speed locomotion requires that braking and propulsion of the stance limbs be equal in magnitude. We predicted that changes in body mass distribution would alter individual limb braking-propulsive force patterns and we tested this hypothesis by adding 10% body mass near the center of mass, at the pectoral girdle, or at the pelvic girdle of trotting dogs. Two force platforms in series recorded fore- and hindlimb ground reaction forces independently. Vertical and fore-aft impulses were calculated by integrating individual force-time curves and Fourier analysis was used to quantify the braking-propulsive (b-p) bias of the fore-aft force curve. We predicted that experimental manipulation of antero-posterior mass distribution would (1) change the fore-hind distribution of vertical impulse when the limb girdles are loaded, (2) decrease the b-p bias of the experimentally loaded limb and (3) increase relative contact time of the experimentally loaded limb, while (4) the individual limb mean fore-aft forces (normalized to body weight + added weight) would be unaffected. All four of these results were observed when mass was added at the pelvic girdle, but only 1, 3 and 4 were observed when mass was added at the pectoral girdle. We propose that the observed relationship between antero-posterior mass distribution and individual limb function may be broadly applicable to quadrupeds with different body types. In addition to the predicted results, our data show that the mechanical effects of adding mass to the trunk are much more complex than would be predicted from mass distribution alone. Effects of trunk moments due to loading were evident when mass was added at the center of mass or at the pelvic girdle. These results suggest a functional link between appendicular and axial mechanics via action of the limbs as levers.

Analysis of Variance↗
PubMed2002

Genetic basis for systems of skeletal quantitative traits: principal component analysis of the canid skeleton.

Evolution of mammalian skeletal structure can be rapid and the changes profound, as illustrated by the morphological diversity of the domestic dog. Here we use principal component analysis of skeletal variation in a population of Portuguese Water Dogs to reveal systems of traits defining skeletal structures. This analysis classifies phenotypic variation into independent components that can be used to dissect genetic networks regulating complex biological systems. We show that unlinked quantitative trait loci associated with these principal components individually promote both correlations within structures (e.g., within the skull or among the limb bones) and inverse correlations between structures (e.g., skull vs. limb bones). These quantitative trait loci are consistent with regulatory genes that inhibit growth of some bones while enhancing growth of others. These systems of traits could explain the skeletal differences between divergent breeds such as Greyhounds and Pit Bulls, and even some of the skeletal transformations that characterize the evolution of hominids.

Animals↗
PubMed2002

The face that sank the Essex: potential function of the spermaceti organ in aggression.

'Forehead to forehead I meet thee, this third time, Moby Dick!' [Ahab (Melville, 1851)] Herman Melville's fictional portrayal of the sinking of the Pequod was inspired by instances in which large sperm whales sank whaling ships by ramming the ships with their heads. Observations of aggression in species of the four major clades of cetacean and the artiodactyl outgroup suggest that head-butting during male-male aggression is a basal behavior for cetaceans. We hypothesize that the ability of sperm whales to destroy stout wooden ships, 3-5 times their body mass, is a product of specialization for male-male aggression. Specifically, we suggest that the greatly enlarged and derived melon of sperm whales, the spermaceti organ, evolved as a battering ram to injure an opponent. To address this hypothesis, we examined the correlation between relative melon size and the level of sexual dimorphism in body size among cetaceans. We also modeled impacts between two equal-sized sperm whales to determine whether it is physically possible for the spermaceti organ to function as an effective battering ram. We found (i) that the evolution of relative melon size in cetaceans is positively correlated with the evolution of sexual dimorphism in body size and (ii) that the spermaceti organ of a charging sperm whale has enough momentum to seriously injure an opponent. These observations are consistent with the hypothesis that the spermaceti organ has evolved to be a weapon used in male-male aggression.

Aggression↗
PubMed2002

Hypaxial muscle activity during running and breathing in dogs.

The axial muscles of terrestrial vertebrates serve two potentially conflicting functions, locomotion and lung ventilation. To differentiate the locomotor and ventilatory functions of the hypaxial muscles in mammals, we examined the locomotor and ventilatory activity of the trunk muscles of trotting dogs under two conditions: when the ventilatory cycle and the locomotor cycle were coupled and when they were uncoupled. Patterns of muscle-activity entrainment with locomotor and ventilatory events revealed (i) that the internal and external abdominal oblique muscles performed primarily locomotor functions during running yet their activity was entrained to expiration when the dogs were standing, (ii) that the internal and external intercostal, external oblique thoracic and transversus abdominis muscles performed both locomotor and respiratory functions simultaneously, (iii) that the parasternal internal intercostal muscle performed a primarily respiratory function (inspiration) and (iv) that the deep pectoralis and longissimus dorsi muscles performed only locomotor functions and were not active while the dogs were standing still. We conclude that the dual function of many hypaxial muscles may produce functional conflicts during running. The redundancy and complexity of the respiratory musculature as well as the particular pattern of respiratory-locomotor coupling in quadrupedal mammals may circumvent these conflicts or minimize their impact on respiration.

Animals↗
PubMed2002

Scaling of rotational inertia in murine rodents and two species of lizard.

Because the force required to rotate a body about an axis is directly proportional to its rotational inertia about the axis, it is likely that animals with high rotational inertia would be constrained in their turning abilities. Given that rotational inertia scales with mass(1.67) in geometrically similar animals, whereas the ability to apply torque scales with mass(1.00), larger animals would be expected to have more difficulty turning than smaller animals of similar shape. To determine how rotational inertia scales with body mass, we used the fact that the period of a physical pendulum is proportional to its rotational inertia(0.50), and measured rotational inertia in two groups of vertebrates with greatly different body shapes: murine rodents (Mus domesticus and Rattus norvegicus) and lizards (Iguana iguana and Varanus exanthematicus). Rotational inertia did not deviate significantly from isometric scaling in the murine rodents as a group or in the varanid lizards, scaling with mass(1.63) and mass(1.59), respectively. Although rotational inertia did scale with negative allometry in iguanas and rats alone, with mass(1.56) and mass(1.42), respectively, it still increased much more quickly with increasing mass than the predicted ability to apply torque. This suggests either that these animals are not constrained by rotational inertia because of their relatively small size or that larger rodents and lizards are poorer turners than smaller ones. The murine rodents had a 3.0- to 4.9-fold lower rotational inertia than similarly sized lizards of either species. Given that the basal synapsids had body proportions and limb configurations similar to those of modern lizards, we suggest that the loss of the large muscular tail and elongated body form during the evolution of cynodonts and mammals reduced rotational inertia and probably improved turning ability.

Animals↗
PubMedDate not supplied

Evidence for endothermic ancestors of crocodiles at the stem of archosaur evolution.

Physiological, anatomical, and developmental features of the crocodilian heart support the paleontological evidence that the ancestors of living crocodilians were active and endothermic, but the lineage reverted to ectothermy when it invaded the aquatic, ambush predator niche. In endotherms, there is a functional nexus between high metabolic rates, high blood flow rates, and complete separation of high systemic blood pressure from low pulmonary blood pressure in a four-chambered heart. Ectotherms generally lack all of these characteristics, but crocodilians retain a four-chambered heart. However, crocodilians have a neurally controlled, pulmonary bypass shunt that is functional in diving. Shunting occurs outside of the heart and involves the left aortic arch that originates from the right ventricle, the foramen of Panizza between the left and right aortic arches, and the cog-tooth valve at the base of the pulmonary artery. Developmental studies show that all of these uniquely crocodilian features are secondarily derived, indicating a shift from the complete separation of blood flow of endotherms to the controlled shunting of ectotherms. We present other evidence for endothermy in stem archosaurs and suggest that some dinosaurs may have inherited the trait.

Alligators and Crocodiles↗