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1.  Directional Asymmetry in the Limbs, Skull and Pelvis of the Silver Fox (V. vulpes) 
Journal of morphology  2010;271(12):1501-1508.
Directional asymmetry (DA) is a characteristic of most vertebrates, most strikingly exhibited by the placement of various organs (heart, lungs, liver, etc.) but also noted in small differences in the metrics of skeletal structures such as the pelvis of certain fish or sauropsids. We have analyzed DA in the skeleton of the fox (V. vulpes), using ~1,000 radiographs of foxes from populations used in the genetic analysis of behavior and morphology. Careful measurements from this robust data base demonstrate that: 1) DA occurs in the limb bones, the ileum, and ischium and in the mandible; 2) regardless of the direction of the length asymmetry vector of a particular skeletal unit, the vectorial direction of length is always opposite to that of width; 3) with the exception of the humerus and radius, there is no correlation or inverse correlation between vectorial amplitudes or magnitudes of bone asymmetries. 4) Postnatal measurements on foxes demonstrate that the asymmetry increases after birth and continues to change (increasing or decreasing) during postnatal growth. 5) A behavior test for preferential use of a specific forelimb exhibited fluctuating asymmetry but not DA. None of the skeletal asymmetries were significantly correlated with a preferential use of a specific forelimb. We suggest that for the majority of fox skeletal parameters, growth on the right and left side of the fox are differentially biased resulting in fixed differences between the two sides in either the rate of growth or the length of the period during which growth occurs. Random effects around these fixed differences perturb the magnitude of the effects such that the magnitudes of length and width asymmetries are not inversely correlated at the level of individual animals.
doi:10.1002/jmor.10890
PMCID: PMC3057660  PMID: 20862692
fox; V. vulpes; skeleton; directional asymmetry; pelvis; mandible; limb bone
2.  A three-dimensional analysis of the morphological evolution and locomotor behaviour of the carnivoran hind limb 
Background
The shape of the appendicular bones in mammals usually reflects adaptations towards different locomotor abilities. However, other aspects such as body size and phylogeny also play an important role in shaping bone design.
We used 3D landmark-based geometric morphometrics to analyse the shape of the hind limb bones (i.e., femur, tibia, and pelvic girdle bones) of living and extinct terrestrial carnivorans (Mammalia, Carnivora) to quantitatively investigate the influence of body size, phylogeny, and locomotor behaviour in shaping the morphology of these bones. We also investigated the main patterns of morphological variation within a phylogenetic context.
Results
Size and phylogeny strongly influence the shape of the hind limb bones. In contrast, adaptations towards different modes of locomotion seem to have little influence. Principal Components Analysis and the study of phylomorphospaces suggest that the main source of variation in bone shape is a gradient of slenderness-robustness.
Conclusion
The shape of the hind limb bones is strongly influenced by body size and phylogeny, but not to a similar degree by locomotor behaviour. The slender-robust “morphological bipolarity” found in bone shape variability is probably related to a trade-off between maintaining energetic efficiency and withstanding resistance to stresses. The balance involved in this trade-off impedes the evolution of high phenotypic variability. In fact, both morphological extremes (slender/robust) are adaptive in different selective contexts and lead to a convergence in shape among taxa with extremely different ecologies but with similar biomechanical demands. Strikingly, this “one-to-many mapping” pattern of evolution between morphology and ecology in hind limb bones is in complete contrast to the “many-to-one mapping” pattern found in the evolution of carnivoran skull shape. The results suggest that there are more constraints in the evolution of the shape of the appendicular skeleton than in that of skull shape because of the strong biomechanical constraints imposed by terrestrial locomotion.
doi:10.1186/1471-2148-14-129
PMCID: PMC4065579  PMID: 24927753
Carnivora; Hind limb; Allometry; Locomotion; Phenotypic evolution; Convergence
3.  Molecular and histopathological detection of Hepatozoon canis in red foxes (Vulpes vulpes) from Portugal 
Parasites & Vectors  2014;7:113.
Background
Hepatozoon canis is a protozoan tick-borne pathogen of dogs and wild canids. Hepatozoon spp. have been reported to infect foxes in different continents and recent studies have mostly used the polymerase chain reaction (PCR) for the detection and characterization of the infecting species. Surveying red foxes (Vulpes vulpes) may contribute to better understanding the epidemiology of canine vector-borne diseases, including hepatozoonosis caused by H. canis in domestic dogs. The present study investigated the prevalence of Hepatozoon spp. by means of histopathology and molecular analysis of different tissues in red foxes from different parts of Portugal.
Methods
Blood and tissues including bone marrow, heart, hind leg muscle, jejunum, kidney, liver, lung, popliteal or axillary lymph nodes, spleen and/or tongue were collected from 91 red foxes from eight districts in northern, central and southern Portugal. Tissues were formalin-fixed, paraffin-embedded, cut and stained with hematoxylin and eosin. Polymerase chain reaction (PCR) amplified a ~650 bp fragment of the 18S rRNA gene of Hepatozoon spp. and the DNA products were sequenced.
Results
Hepatozoon canis was detected in 68 out of 90 foxes (75.6%) from all the sampled areas by PCR and sequencing. Histopathology revealed H. canis meronts similar in shape to those found in dogs in the bone marrow of 11 (23.4%) and in the spleen of two (4.3%) out of 47 foxes (p = 0.007). All the 11 foxes found positive by histopathology were also positive by PCR of bone marrow and/or blood. Positivity by PCR (83.0%) was significantly higher (p < 0.001) than by histopathological examination (23.4%) in paired bone marrow samples from the same 47 foxes. Sequences of the 18S rRNA gene of H. canis were 98–99% identical to those in GenBank.
Conclusions
Hepatozoon canis was found to be highly prevalent in red fox populations from northern, central and southern Portugal. Detection of the parasite by histopathology was significantly less sensitive than by PCR. Red foxes are a presumptive reservoir of H. canis infection for domestic dogs.
doi:10.1186/1756-3305-7-113
PMCID: PMC3994325  PMID: 24655375
Canine vector-borne diseases; Hepatozoon canis; Histopathology; Polymerase chain reaction; Portugal; Red fox; Vulpes vulpes
4.  Radiographic analysis of vocal tract length and its relation to overall body size in two canid species 
Journal of zoology (London, England : 1987)  2013;291(1):10.1111/jzo.12048.
Body size is an important determinant of resource and mate competition in many species. Competition is often mediated by conspicuous vocal displays, which may help to intimidate rivals and attract mates by providing honest cues to signaler size. Fitch proposed that vocal tract resonances (or formants) should provide particularly good, or honest, acoustic cues to signaler size because they are determined by the length of the vocal tract, which in turn, is hypothesized to scale reliably with overall body size. There is some empirical support for this hypothesis, but to date, many of the effects have been either mixed for males compared with females, weaker than expected in one or the other sex, or complicated by sampling issues. In this paper, we undertake a direct test of Fitch’s hypothesis in two canid species using large samples that control for age- and sex-related variation. The samples involved radiographic images of 120 Portuguese water dogs Canis lupus familiaris and 121 Russian silver foxes Vulpes vulpes. Direct measurements were made of vocal tract length from X-ray images and compared against independent measures of body size. In adults of both species, and within both sexes, overall vocal tract length was strongly and significantly correlated with body size. Effects were strongest for the oral component of the vocal tract. By contrast, the length of the pharyngeal component was not as consistently related to body size. These outcomes are some of the clearest evidence to date in support of Fitch’s hypothesis. At the same time, they highlight the potential for elements of both honest and deceptive body signaling to occur simultaneously via differential acoustic cues provided by the oral versus pharyngeal components of the vocal tract.
doi:10.1111/jzo.12048
PMCID: PMC3866914  PMID: 24363497
body size; vocal tract length; animal communication
5.  A Three-Dimensional Analysis of Morphological Evolution and Locomotor Performance of the Carnivoran Forelimb 
PLoS ONE  2014;9(1):e85574.
In this study, three-dimensional landmark-based methods of geometric morphometrics are used for estimating the influence of phylogeny, allometry and locomotor performance on forelimb shape in living and extinct carnivorans (Mammalia, Carnivora). The main objective is to investigate morphological convergences towards similar locomotor strategies in the shape of the major forelimb bones. Results indicate that both size and phylogeny have strong effects on the anatomy of all forelimb bones. In contrast, bone shape does not correlate in the living taxa with maximum running speed or daily movement distance, two proxies closely related to locomotor performance. A phylomorphospace approach showed that shape variation in forelimb bones mainly relates to changes in bone robustness. This indicates the presence of biomechanical constraints resulting from opposite demands for energetic efficiency in locomotion –which would require a slender forelimb– and resistance to stress –which would be satisfied by a robust forelimb–. Thus, we interpret that the need of maintaining a trade-off between both functional demands would limit shape variability in forelimb bones. Given that different situations can lead to one or another morphological solution, depending on the specific ecology of taxa, the evolution of forelimb morphology represents a remarkable “one-to-many mapping” case between anatomy and ecology.
doi:10.1371/journal.pone.0085574
PMCID: PMC3893248  PMID: 24454891
6.  Anatomical Contributions to Hylobatid Taxonomy and Adaptation 
Compared with the great apes, the small-bodied hylobatids were treated historically as a relatively uniform group with 2 genera, Hylobates and the larger-bodied Symphalangus. Four genera are now recognized, each with a different chromosome number: Hoolock (hoolock) (38), Hylobates (44), Nomascus (crested gibbon) (52), and Symphalangus (siamang) (50). Previous morphological studies based on relative bone lengths, e.g., intermembral indices; molar tooth sizes; and body masses did not distinguish the 4 genera from each other. We applied quantitative anatomical methods to test the hypothesis that each genus can be differentiated from the others using the relative distribution of body mass to the forelimbs and hind limbs. Based on dissections of 13 hylobatids from captive facilities, our findings demonstrate that each of the 4 genera has a distinct pattern of body mass distribution. For example, the adult Hoolock has limb proportions of nearly equal mass, a pattern that differentiates it from species in the genus Hylobates, e.g., H. lar (lar gibbon), H. moloch (Javan gibbon), H. pileatus (pileated gibbon), Nomascus, and Symphalangus. Hylobates is distinct in having heavy hind limbs. Although Symphalangus has been treated as a scaled up version of Hylobates, its forelimb exceeds its hind limb mass, an unusual primate pattern otherwise found only in orangutans. This research provides new information on whole body anatomy and adds to the genetic, ecological, and behavioral evidence for clarifying the taxonomy of the hylobatids. The research also underscores the important contribution of studies on rare species in captivity.
doi:10.1007/s10764-011-9506-y
PMCID: PMC3139862  PMID: 21892236
Anatomy; Body proportions; Hylobates; Hylobatids; Limb mass; Symphalangus; Gibbon; Siamang; Taxonomy
7.  Chromosomal Mapping of Canine-Derived BAC Clones to the Red Fox and American Mink Genomes 
Journal of Heredity  2009;100(Suppl 1):S42-S53.
High-quality sequencing of the dog (Canis lupus familiaris) genome has enabled enormous progress in genetic mapping of canine phenotypic variation. The red fox (Vulpes vulpes), another canid species, also exhibits a wide range of variation in coat color, morphology, and behavior. Although the fox genome has not yet been sequenced, canine genomic resources have been used to construct a meiotic linkage map of the red fox genome and begin genetic mapping in foxes. However, a more detailed gene-specific comparative map between the dog and fox genomes is required to establish gene order within homologous regions of dog and fox chromosomes and to refine breakpoints between homologous chromosomes of the 2 species. In the current study, we tested whether canine-derived gene–containing bacterial artificial chromosome (BAC) clones can be routinely used to build a gene-specific map of the red fox genome. Forty canine BAC clones were mapped to the red fox genome by fluorescence in situ hybridization (FISH). Each clone was uniquely assigned to a single fox chromosome, and the locations of 38 clones agreed with cytogenetic predictions. These results clearly demonstrate the utility of FISH mapping for construction of a whole-genome gene-specific map of the red fox. The further possibility of using canine BAC clones to map genes in the American mink (Mustela vison) genome was also explored. Much lower success was obtained for this more distantly related farm-bred species, although a few BAC clones were mapped to the predicted chromosomal locations.
doi:10.1093/jhered/esp037
PMCID: PMC3139363  PMID: 19546120
Canis lupus familiaris; comparative genomics; FISH; Mustela vison; Vulpes vulpes
8.  Evolution of adaptive phenotypic variation patterns by direct selection for evolvability 
A basic assumption of the Darwinian theory of evolution is that heritable variation arises randomly. In this context, randomness means that mutations arise irrespective of the current adaptive needs imposed by the environment. It is broadly accepted, however, that phenotypic variation is not uniformly distributed among phenotypic traits, some traits tend to covary, while others vary independently, and again others barely vary at all. Furthermore, it is well established that patterns of trait variation differ among species. Specifically, traits that serve different functions tend to be less correlated, as for instance forelimbs and hind limbs in bats and humans, compared with the limbs of quadrupedal mammals. Recently, a novel class of genetic elements has been identified in mouse gene-mapping studies that modify correlations among quantitative traits. These loci are called relationship loci, or relationship Quantitative Trait Loci (rQTL), and affect trait correlations by changing the expression of the existing genetic variation through gene interaction. Here, we present a population genetic model of how natural selection acts on rQTL. Contrary to the usual neo-Darwinian theory, in this model, new heritable phenotypic variation is produced along the selected dimension in response to directional selection. The results predict that selection on rQTL leads to higher correlations among traits that are simultaneously under directional selection. On the other hand, traits that are not simultaneously under directional selection are predicted to evolve lower correlations. These results and the previously demonstrated existence of rQTL variation, show a mechanism by which natural selection can directly enhance the evolvability of complex organisms along lines of adaptive change.
doi:10.1098/rspb.2010.2113
PMCID: PMC3097830  PMID: 21106581
modularity; allometry; canalization; differential epistasis
9.  Temporospatial and kinetic characteristics of sheep walking on a pressure sensing walkway 
The walking gait of sheep was analyzed in terms of temporospatial and kinetic parameters and weight distribution among the 4 limbs. Eighteen mature female Suffolk-mix sheep walked comfortably with a halter-guide over a 1.5-m pressure sensing walkway. Six valid trials were acquired for each sheep without any previous selection or habituation. Stance phases of the forelimb and hind limb were 66.3% and 68.9%, respectively, of total gait cycle, and limb velocity was 1.06 m/s in both forelimbs and hind limbs while walking. The mean peak vertical force (PVF) and vertical impulse (VI) as percentage of body weight in the forelimbs were 52.5% and 19.9%, respectively, and those of the hind limbs were 38.5% and 14.9%, respectively. More body weight was loaded on the forelimbs than the hind limbs, at 59% and 41% of body weight, respectively. The walking gait of sheep measured with the pressure sensing (PS) walkway was similar to that reported in dogs and horses. The PS walkway enabled collection of temporospatial and kinetic data, and simplified the process of data collection.
PMCID: PMC2117367  PMID: 18214162
10.  Kinematic analysis of forelimb and hind limb joints in clinically healthy sheep 
BMC Veterinary Research  2014;10(1):294.
Background
Variations associated with sex, age, velocity, breed and body geometry should be considered in the determination of kinematic parameters for a gait considered normal.
Therefore, this study aimed to evaluate kinematic patterns of forelimbs and hind limbs in clinically normal sheep from two different age groups walking at a constant velocity. The hypothesis was that the age may influence sagittal plane kinematic patterns. Fourteen clinically healthy female sheep were divided into Group 1 – seven animals aged from 8 to 12 months, and Group 2 - seven animals aged above 5 years. Before starting data collection, the sheep were trained to be conducted for walking in a pre-determined space at constant velocity. A minimum of 5 valid trials were obtained from the right and left sides of each sheep. Data were analyzed by use of a motion-analysis program. Flexion and extension joint angles (maximum, minimum, displacement), and angular velocity (maximum, minimum) were determined for the shoulder, elbow, carpal, hip, stifle, and tarsal joints.
Results
Within each group, no significant differences were observed between the right and left limbs in all kinematic variables. Significant differences were observed in the following kinematic parameters between G1 and G2: minimum angle (G1 < G2), angular displacement (G1 > G2), maximum angular velocity (G1 > G2), minimum angular velocity (G1 > G2) of the carpus; angular displacement (G1 > G2), minimum angular velocity (G1 > G2) of the shoulder; minimum angle (G1 > G2), angular displacement (G1 < G2) of the tarsus; maximum angular velocity (G1 < G2) of the stifle; maximum angular velocity G1 > G2 of the hip. The lengths of both forelimbs and hind limbs differed between groups (G1 < G2). The Froude number differed between groups for forelimbs and hind limbs.
Conclusions
In conclusion, sheep of two different ages walking at a constant velocity present, within the same group, similar kinematic data between sides, and exhibit some differences in kinematic variables that may be age-related or body size. Further studies using sheep walking at similar Froude numbers are necessary to exclude the body size.
doi:10.1186/s12917-014-0294-4
PMCID: PMC4269177  PMID: 25495531
11.  Heritabilities of Directional Asymmetry in the Fore- and Hindlimbs of Rabbit Fetuses 
PLoS ONE  2013;8(10):e76358.
Directional asymmetry (DA), where at the population level symmetry differs from zero, has been reported in a wide range of traits and taxa, even for traits in which symmetry is expected to be the target of selection such as limbs or wings. In invertebrates, DA has been suggested to be non-adaptive. In vertebrates, there has been a wealth of research linking morphological asymmetry to behavioural lateralisation. On the other hand, the prenatal expression of DA and evidences for quantitative genetic variation for asymmetry may suggest it is not solely induced by differences in mechanic loading between sides. We estimate quantitative genetic variation of fetal limb asymmetry in a large dataset of rabbits. Our results showed a low but highly significant level of DA that is partially under genetic control for all traits, with forelimbs displaying higher levels of asymmetry. Genetic correlations were positive within limbs, but negative across bones of fore and hind limbs. Environmental correlations were positive for all, but smaller across fore and hind limbs. We discuss our results in light of the existence and maintenance of DA in locomotory traits.
doi:10.1371/journal.pone.0076358
PMCID: PMC3794934  PMID: 24130770
12.  Ontogenetic allometry of the Beagle 
Background
Mammalian juveniles undergo dramatic changes in body conformation during development. As one of the most common companion animals, the time line and trajectory of a dog’s development and its body’s re-proportioning is of particular scientific interest. Several ontogenetic studies have investigated the skeletal development in dogs, but none has paid heed to the scapula as a critical part of the mammalian forelimb. Its functional integration into the forelimb changed the correspondence between fore- and hindlimb segments and previous ontogenetic studies observed more similar growth patterns for functionally than serially homologous elements. In this study, the ontogenetic development of six Beagle siblings was monitored between 9 and 51 weeks of age to investigate their skeletal allometry and compare this with data from other lines, breeds and species.
Results
Body mass increased exponentially with time; log linear increase was observed up to the age of 15 weeks. Compared with body mass, withers and pelvic height as well as the lengths of the trunk, scapula, brachium and antebrachium, femur and crus exhibited positive allometry. Trunk circumference and pes showed negative allometry in all, pelvis and manus in most dogs. Thus, the typical mammalian intralimb re-proportioning with the proximal limb elements exhibiting positive allometry and the very distal ones showing negative allometry was observed. Relative lengths of the antebrachium, femur and crus increased, while those of the distal elements decreased.
Conclusions
Beagles are fully-grown regarding body height but not body mass at about one year of age. Particular attention should be paid to feeding and physical exertion during the first 15 weeks when they grow more intensively. Compared with its siblings, a puppy’s size at 9 weeks is a good indicator for its final size. Among siblings, growth duration may vary substantially and appears not to be related to the adult size. Within breeds, a longer time to physically mature is hypothesized for larger-bodied breeding lines. Similar to other mammals, the Beagle displayed nearly optimal intralimb proportions throughout development. Neither the forelimbs nor the hindlimbs conformed with the previously observed proximo-distal order of the limb segment’s growth gradients. Potential factors responsible for variations in the ontogenetic allometry of mammals need further evaluation.
doi:10.1186/1746-6148-9-203
PMCID: PMC4124762  PMID: 24112520
Scaling; Limb proportions; Body proportions; Bone growth; Serial homology; Body mass
13.  Ontogenetic Scaling of Fore- and Hind Limb Posture in Wild Chacma Baboons (Papio hamadryas ursinus) 
PLoS ONE  2013;8(7):e71020.
Large-scale interspecific studies of mammals ranging between 0.04–280 kg have shown that larger animals walk with more extended limb joints. Within a taxon or clade, however, the relationship between body size and joint posture is less straightforward. Factors that may affect the lack of congruence between broad and narrow phylogenetic analyses of limb kinematics include limited sampling of (1) ranges of body size, and/or (2) numbers of individuals. Unfortunately, both issues are inherent in laboratory-based or zoo locomotion research. In this study, we examined the relationship between body mass and elbow and knee joint angles (our proxies of fore- and hind limb posture, respectively) in a cross-sectional ontogenetic sample of wild chacma baboons (Papio hamadryas ursinus) habituated in the De Hoop Nature Reserve, South Africa. Videos were obtained from 33 individuals of known age (12 to ≥108 months) and body mass (2–29.5 kg) during walking trials. Results show that older, heavier baboons walk with significantly more extended knee joints but not elbow joints. This pattern is consistent when examining only males, but not within the female sample. Heavier, older baboons also display significantly less variation in their hind limb posture compared to lighter, young animals. Thus, within this ontogenetic sample of a single primate species spanning an order of magnitude in body mass, hind limb posture exhibited a postural scaling phenomenon while the forelimbs did not. These findings may further help explain 1) why younger mammals (including baboons) tend to have relatively stronger bones than adults, and 2) why humeri appear relatively weaker than femora (in at least baboons). Finally, this study demonstrates how field-acquired kinematics can help answer fundamental biomechanical questions usually addressed only in animal gait laboratories.
doi:10.1371/journal.pone.0071020
PMCID: PMC3726614  PMID: 23923046
14.  A Simple Genetic Architecture Underlies Morphological Variation in Dogs 
PLoS Biology  2010;8(8):e1000451.
The largest genetic study to date of morphology in domestic dogs identifies genes controlling nearly 100 morphological traits and identifies important trends in phenotypic variation within this species.
Domestic dogs exhibit tremendous phenotypic diversity, including a greater variation in body size than any other terrestrial mammal. Here, we generate a high density map of canine genetic variation by genotyping 915 dogs from 80 domestic dog breeds, 83 wild canids, and 10 outbred African shelter dogs across 60,968 single-nucleotide polymorphisms (SNPs). Coupling this genomic resource with external measurements from breed standards and individuals as well as skeletal measurements from museum specimens, we identify 51 regions of the dog genome associated with phenotypic variation among breeds in 57 traits. The complex traits include average breed body size and external body dimensions and cranial, dental, and long bone shape and size with and without allometric scaling. In contrast to the results from association mapping of quantitative traits in humans and domesticated plants, we find that across dog breeds, a small number of quantitative trait loci (≤3) explain the majority of phenotypic variation for most of the traits we studied. In addition, many genomic regions show signatures of recent selection, with most of the highly differentiated regions being associated with breed-defining traits such as body size, coat characteristics, and ear floppiness. Our results demonstrate the efficacy of mapping multiple traits in the domestic dog using a database of genotyped individuals and highlight the important role human-directed selection has played in altering the genetic architecture of key traits in this important species.
Author Summary
Dogs offer a unique system for the study of genes controlling morphology. DNA from 915 dogs from 80 domestic breeds, as well as a set of feral dogs, was tested at over 60,000 points of variation and the dataset analyzed using novel methods to find loci regulating body size, head shape, leg length, ear position, and a host of other traits. Because each dog breed has undergone strong selection by breeders to have a particular appearance, there is a strong footprint of selection in regions of the genome that are important for controlling traits that define each breed. These analyses identified new regions of the genome, or loci, that are important in controlling body size and shape. Our results, which feature the largest number of domestic dogs studied at such a high level of genetic detail, demonstrate the power of the dog as a model for finding genes that control the body plan of mammals. Further, we show that the remarkable diversity of form in the dog, in contrast to some other species studied to date, appears to have a simple genetic basis dominated by genes of major effect.
doi:10.1371/journal.pbio.1000451
PMCID: PMC2919785  PMID: 20711490
15.  Kinetic and temporospatial parameters in male and female cats walking over a pressure sensing walkway 
Background
Several factors may influence kinetic data measurements, including body conformation and body mass. In addition, gender differences in gait pattern have been observed in healthy humans. Therefore, the aim of this study was to compare the kinetic and temporospatial parameters in clinically healthy male and female cats using a pressure-sensitive walkway. Eighteen crossbreed adult cats were divided into two groups: G1 had ten male cats (nine neutered) aged from 1 to 4 years and body mass 3.1-6.8 kg; G2 had eight spayed female cats, aged from 1 to 6 years and body mass 3.3-4.75 kg. The data from the first five valid trials were collected for each cat. A trial was considered valid if the cat maintained a velocity between 0.54-0.74 m/s and acceleration from -0.20 to 0.20 m/s2. The peak vertical force (PVF), vertical impulse (VI), gait cycle time, stance time, swing time, stride length, and percentage body weight distribution among the four limbs were determined. In addition, the lengths of each forelimb and each hind limb were measured using a tape with the animal standing.
Results
No significant differences were observed in each group in either the forelimbs or the hind limbs or between the left and right sides for any of the variables. For both groups, the PVF (%BW), the VI, and the percentage body weight distribution were higher at the forelimbs than the hind limbs. The stride length was larger for males; however, the other kinetic and temporospatial variables did not show any statistically significant differences between the groups. The lengths of the forelimbs and hind limbs were larger in the male cats. There was a significant moderate positive correlation between the stride length and the length of the limbs.
Conclusions
In conclusion, the only difference observed between male and female cats was the stride length, and this was due to the greater body size of male cats. This difference did not affect other temporospatial or kinetics variables.
doi:10.1186/1746-6148-9-129
PMCID: PMC3701551  PMID: 23803220
16.  Pressure mat analysis of the longitudinal development of pig locomotion in growing pigs after weaning 
Background
Gait evaluation is difficult in pigs, especially when objective and quantitative data are needed, thus little research has been conducted in this species. There is considerable experience, however, with objective gait analysis in other species, such as horses and dogs. In this study, a pressure mat was used to establish baseline kinetic data for gait and its longitudinal development in growing, weaned piglets.
Ten clinically healthy weaned piglets were trained to trot over a pressure mat. Measurements were performed weekly during 10 weeks, starting at 5 weeks of age. Four kinetic parameters were recorded for all four limbs: peak vertical force (PVF), load rate (LR), vertical impulse (VI) and peak vertical pressure (PVP). Three representative runs per measuring session per pig were collected. For each of the variables, left vs. right limb asymmetry-indices (ASI’s) were calculated based on the average for that parameter per week. A linear mixed model was used to determine the influence of time (week), velocity, and limb (left vs. right, and fore vs. hind). Intra-class correlations were calculated to assess within-session replicability.
Results
Intra-class correlations showed good within-session replicability. Body-weight normalized PVF (nPVF), LR (nLR), VI (nVI) and PVP (nPVP) were higher in the forelimbs than in the hind limbs. A higher velocity was associated with a higher nPVF, nLR and nPVP. All parameters varied between weeks. ASI of LR and VI were higher in the forelimbs than in the hind limbs. Velocity and time did not influence ASI of any of the variables.
Conclusions
Kinetic pressure mat measurements from healthy weaned piglets are highly replicable within-session. However, these variables present a significant variability between-session, which may be due to conformational changes of the young, growing piglets. Velocity clearly influences nPVF, nLR and nPVP, and all kinetic variables have higher values in forelimbs than in hind limbs. As time and velocity do not affect ASI’s, the latter are preferable tools when velocity cannot be controlled or when measurements are repeated over longer time intervals. The present study supports the use of a pressure mat as an objective way to analyze and quantify porcine gait.
doi:10.1186/1746-6148-10-37
PMCID: PMC3925128  PMID: 24502522
Kinetics; Gait analysis; Pig; Symmetry
17.  Gait analysis in clinically healthy sheep from three different age groups using a pressure-sensitive walkway 
Background
Understanding normal gait requires allowing for variations in normal patterns by the sex, age, and species in question. Therefore, the aim of this study was to evaluate kinetic and temporospatial parameters in clinically healthy sheep from three different age groups with a pressure-sensing walkway. The sheep were judged to be healthy based on the results of complete physical and orthopaedic examinations and had no history of lameness. Twenty-one clinically healthy female Santa Ines sheep were divided into three groups: G1 – seven animals, aged from 8 to 12 months and weighing 19.5-33 kg; G2 - seven individuals, aged from 2 to 4 years and weighing 26.5-42 kg; and G3 - seven sheep, aged more than 5 years and weighing 37.3-45 kg. The animals were examined from two directions: first on the left side and then on the right side of the handler. The data from the first five valid trials in each direction were collected for each sheep and analysed using the designated software. A trial was considered valid if the sheep walked within the correct velocity (1.1-1.3 m/s) and acceleration (from −0.15 to 0.15 m/s2) ranges. The peak vertical force (PVF), vertical impulse (VI), gait cycle time, stance time, swing time, stride length, and the percentage body weight distribution among the four limbs were determined.
Results
No significant differences were observed, in either the forelimbs or the hind limbs, between the left and right sides or between the two directions for any of the variables. No significant temporospatial differences were found among the groups. Significant PVF (%BW) differences were observed in the forelimbs (G1 > G3) and hind limbs (G1 > G3), and significant VI differences were observed in the forelimbs (G1 > G3).
Conclusions
Young healthy sheep differ from older sheep in the vertical forces they create when walking at the same velocity on a pressure-sensing walkway.
doi:10.1186/1746-6148-8-87
PMCID: PMC3418207  PMID: 22726641
18.  Forelimb-hindlimb developmental timing changes across tetrapod phylogeny 
Background
Tetrapods exhibit great diversity in limb structures among species and also between forelimbs and hindlimbs within species, diversity which frequently correlates with locomotor modes and life history. We aim to examine the potential relation of changes in developmental timing (heterochrony) to the origin of limb morphological diversity in an explicit comparative and quantitative framework. In particular, we studied the relative time sequence of development of the forelimbs versus the hindlimbs in 138 embryos of 14 tetrapod species spanning a diverse taxonomic, ecomorphological and life-history breadth. Whole-mounts and histological sections were used to code the appearance of 10 developmental events comprising landmarks of development from the early bud stage to late chondrogenesis in the forelimb and the corresponding serial homologues in the hindlimb.
Results
An overall pattern of change across tetrapods can be discerned and appears to be relatively clade-specific. In the primitive condition, as seen in Chondrichthyes and Osteichthyes, the forelimb/pectoral fin develops earlier than the hindlimb/pelvic fin. This pattern is either retained or re-evolved in eulipotyphlan insectivores (= shrews, moles, hedgehogs, and solenodons) and taken to its extreme in marsupials. Although exceptions are known, the two anurans we examined reversed the pattern and displayed a significant advance in hindlimb development. All other species examined, including a bat with its greatly enlarged forelimbs modified as wings in the adult, showed near synchrony in the development of the fore and hindlimbs.
Conclusion
Major heterochronic changes in early limb development and chondrogenesis were absent within major clades except Lissamphibia, and their presence across vertebrate phylogeny are not easily correlated with adaptive phenomena related to morphological differences in the adult fore- and hindlimbs. The apparently conservative nature of this trait means that changes in chondrogenetic patterns may serve as useful phylogenetic characters at higher taxonomic levels in tetrapods. Our results highlight the more important role generally played by allometric heterochrony in this instance to shape adult morphology.
doi:10.1186/1471-2148-7-182
PMCID: PMC2194785  PMID: 17908305
19.  Quantitative Phenotyping of Duchenne Muscular Dystrophy Dogs by Comprehensive Gait Analysis and Overnight Activity Monitoring 
PLoS ONE  2013;8(3):e59875.
The dystrophin-deficient dog is excellent large animal model for testing novel therapeutic modalities for Duchenne muscular dystrophy (DMD). Despite well-documented descriptions of dystrophic symptoms in these dogs, very few quantitative studies have been performed. Here, we developed a comprehensive set of non-invasive assays to quantify dog gait (stride length and speed), joint angle and limb mobility (for both forelimb and hind limb), and spontaneous activity at night. To validate these assays, we examined three 8-m-old mix-breed dystrophic dogs. We also included three age-matched siblings as the normal control. High-resolution video recorders were used to digitize dog walking and spontaneous movement at night. Stride speed and length were significantly decreased in affected dogs. The mobility of the limb segments (forearm, front foot, lower thigh, rear foot) and the carpus and hock joints was significantly reduced in dystrophic dogs. There was also a significant reduction of the movement in affected dogs during overnight monitoring. In summary, we have established a comprehensive set of outcome measures for clinical phenotyping of DMD dogs. These non-invasive end points would be valuable in monitoring disease progression and therapeutic efficacy in translational studies in the DMD dog model.
doi:10.1371/journal.pone.0059875
PMCID: PMC3609742  PMID: 23544107
20.  Transcriptional heterochrony in talpid mole autopods 
EvoDevo  2012;3:16.
Background
Talpid moles show many specializations in their adult skeleton linked to fossoriality, including enlarged hands when compared to the feet. Heterochrony in developmental mechanisms is hypothesized to account for morphological evolution in skeletal elements.
Methods
The temporal and spatial distribution of SOX9 expression, which is an early marker of chondrification, is analyzed in autopods of the fossorial Iberian mole Talpa occidentalis, as well as in shrew (Cryptotis parva) and mouse (Mus musculus) for comparison.
Results and discussion
SOX9 expression is advanced in the forelimb compared to the hind limb in the talpid mole. In contrast, in the shrew and the mouse, which do not show fossorial specializations in their autopods, it is synchronous. We provide evidence that transcriptional heterochrony affects the development of talpid autopods, an example of developmental penetrance. We discuss our data in the light of earlier reported pattern heterochrony and later morphological variation in talpid limbs.
Conclusion
Transcriptional heterochrony in SOX9 expression is found in talpid autopods, which is likely to account for pattern heterochrony in chondral limb development as well as size variation in adult fore- and hind limbs.
doi:10.1186/2041-9139-3-16
PMCID: PMC3441920  PMID: 22873211
SOX9 expression; Developmental penetrance; Talpidae
21.  Radiographs Reveal Exceptional Forelimb Strength in the Sabertooth Cat, Smilodon fatalis 
PLoS ONE  2010;5(7):e11412.
Background
The sabertooth cat, Smilodon fatalis, was an enigmatic predator without a true living analog. Their elongate canine teeth were more vulnerable to fracture than those of modern felids, making it imperative for them to immobilize prey with their forelimbs when making a kill. As a result, their need for heavily muscled forelimbs likely exceeded that of modern felids and thus should be reflected in their skeletons. Previous studies on forelimb bones of S. fatalis found them to be relatively robust but did not quantify their ability to withstand loading.
Methodology/Principal Findings
Using radiographs of the sabertooth cat, Smilodon fatalis, 28 extant felid species, and the larger, extinct American lion Panthera atrox, we measured cross-sectional properties of the humerus and femur to provide the first estimates of limb bone strength in bending and torsion. We found that the humeri of Smilodon were reinforced by cortical thickening to a greater degree than those observed in any living felid, or the much larger P. atrox. The femur of Smilodon also was thickened but not beyond the normal variation found in any other felid measured.
Conclusions/Significance
Based on the cross-sectional properties of its humerus, we interpret that Smilodon was a powerful predator that differed from extant felids in its greater ability to subdue prey using the forelimbs. This enhanced forelimb strength was part of an adaptive complex driven by the need to minimize the struggles of prey in order to protect the elongate canines from fracture and position the bite for a quick kill.
doi:10.1371/journal.pone.0011412
PMCID: PMC2896400  PMID: 20625398
22.  A Combination of Activation and Repression by a Colinear Hox Code Controls Forelimb-Restricted Expression of Tbx5 and Reveals Hox Protein Specificity 
PLoS Genetics  2014;10(3):e1004245.
Tight control over gene expression is essential for precision in embryonic development and acquisition of the regulatory elements responsible is the predominant driver for evolution of new structures. Tbx5 and Tbx4, two genes expressed in forelimb and hindlimb-forming regions respectively, play crucial roles in the initiation of limb outgrowth. Evolution of regulatory elements that activate Tbx5 in rostral LPM was essential for the acquisition of forelimbs in vertebrates. We identified such a regulatory element for Tbx5 and demonstrated Hox genes are essential, direct regulators. While the importance of Hox genes in regulating embryonic development is clear, Hox targets and the ways in which each protein executes its specific function are not known. We reveal how nested Hox expression along the rostro-caudal axis restricts Tbx5 expression to forelimb. We demonstrate that Hoxc9, which is expressed in caudal LPM where Tbx5 is not expressed, can form a repressive complex on the Tbx5 forelimb regulatory element. This repressive capacity is limited to Hox proteins expressed in caudal LPM and carried out by two separate protein domains in Hoxc9. Forelimb-restricted expression of Tbx5 and ultimately forelimb formation is therefore achieved through co-option of two characteristics of Hox genes; their colinear expression along the body axis and the functional specificity of different paralogs. Active complexes can be formed by Hox PG proteins present throughout the rostral-caudal LPM while restriction of Tbx5 expression is achieved by superimposing a dominant repressive (Hoxc9) complex that determines the caudal boundary of Tbx5 expression. Our results reveal the regulatory mechanism that ensures emergence of the forelimbs at the correct position along the body. Acquisition of this regulatory element would have been critical for the evolution of limbs in vertebrates and modulation of the factors we have identified can be molecular drivers of the diversity in limb morphology.
Author Summary
The acquisition of limbs during vertebrate evolution was a very successful innovation that enabled this group of species to diversify and colonise land. It has become clear recently that the primary driver behind the evolution of new structures, such as limbs, is the acquisition of novel regulatory elements that control when and where genes are activated rather than the proteins encoded by the genes themselves acquiring novel functions. We have identified the regulatory element from a gene, Tbx5. Activation of Tbx5 in the forelimb-forming region of the developing embryos is essential for forelimbs to form and disruption of human TBX5 causes limb abnormalities. We show that activation of Tbx5 in a restricted territory is achieved through a combination of activation inputs that are present broadly throughout the embryo flank and dominant, repressive inputs present only in more caudal regions of the flank. The sum of these inputs yields restricted activation in the rostral, forelimb-forming flank. Our results explain how the regulatory switches that were harnessed for the acquisition of limbs during evolution operate and how they can be turned off during the evolution of limblessness in species such as the snake.
doi:10.1371/journal.pgen.1004245
PMCID: PMC3961185  PMID: 24651482
23.  Comparison of Metal Concentrations in Bones of Long-Living Mammals 
Biological Trace Element Research  2013;152(2):195-203.
The aim of this study was to compare zinc, copper, lead, cadmium, and mercury concentrations in the bones of long-living mammals—humans (Homo sapiens) and Canidae (dogs Canis familiaris and foxes Vulpes vulpes) from northwestern Poland and to determine the usefulness of Canidae as bioindicators of environmental exposure to metals in humans. Zinc concentrations in cartilage with adjacent compact bone and in spongy bone were highest in foxes (∼120 mg/kg dry weight (dw)) and lowest in dogs (80 mg/kg dw). Copper concentrations in cartilage with adjacent compact bone were greatest in foxes (1.17 mg/kg dw) and smallest in humans (∼0.8 mg/kg dw), while in spongy bone they were greatest in dogs (0.76 mg/kg dw) and lowest in foxes (0.45 mg/kg dw). Lead concentrations in both analyzed materials were highest in dogs (>3 mg/kg dw) and lowest in humans (>0.6 mg/kg dw). Cadmium concentration, also in both the analyzed materials, were highest in foxes (>0.15 mg/kg dw) and lowest in humans (>0.04 mg/kg dw). Mercury concentration in bones was low and did not exceed 0.004 mg/kg dw in all the examined species. The concentrations of essential metals in the bones of the examined long-living mammals were similar. The different concentrations of toxic metals were due to environmental factors. As bone tissues are used in the assessment of the long-term effects of environmental exposure to heavy metals on the human body, ecotoxicological studies on the bones of domesticated and wild long-living mammals, including Canidae, may constitute a significant supplement to this research.
doi:10.1007/s12011-013-9615-x
PMCID: PMC3624004  PMID: 23377610
Human; Dog; Red fox; Bioindicator; Bone tissue; Trace elements
24.  Animal evolution during domestication: the domesticated fox as a model 
Summary
We review the evolution of domestic animals, emphasizing the effect of the earliest steps of domestication on its course. Using the first domesticated species, the dog (Canis familiaris) as an illustration, we describe the evolutionary specificities of the historical domestication, such as the high level and wide range of diversity. We suggest that the process of earliest domestication via unconscious and later conscious selection of human-defined behavioral traits may accelerate phenotypic variations. The review is based on the results of the long-term experiment designed to reproduce early mammalian domestication in the silver fox (Vulpes vulpes) selected for tameability, or amenability to domestication. We describe changes in behavior, morphology and physiology that appeared in the fox during its selection for tameability and that were similar to those observed in the domestic dog. Based on the experimental fox data and survey of relevant data, we discuss the developmental, genetic and possible molecular-genetic mechanisms of these changes. We assign the causative role in evolutionary transformation of domestic animals to selection for behavior and to the neurospecific regulatory genes it affects.
doi:10.1002/bies.200800070
PMCID: PMC2763232  PMID: 19260016
25.  Genetic correlations between conformation traits and radiographic findings in the limbs of German Warmblood riding horses 
Studbook inspection (SBI) data of 20 768 German Warmblood mares and radiography results (RR) data of 5102 Hanoverian Warmblood horses were used for genetic correlation analyses. The scores on a scale from 0 to 10 were given for conformation and basic quality of gaits, resulting in 14 SBI traits which were used for the correlation analyses. The radiographic findings considered included osseous fragments in fetlock (OFF) and hock joints (OFH), deforming arthropathy in hock joints (DAH) and distinct radiographic findings in the navicular bones (DNB) which were analyzed as binary traits, and radiographic appearance of the navicular bones (RNB) which was analyzed as a quasi-linear trait. Genetic parameters were estimated multivariately in linear animal models with REML using information on 24 448 horses with SBI and/or RR records. The ranges of heritability estimates were h2 = 0.14–0.34 for the RR traits and h2 = 0.09–0.50 for the SBI traits. Negative additive genetic correlations of rg = -0.19 to -0.56 were estimated between OFF and conformation of front and hind limbs and walk at hand, and between DNB and hind limb conformation. There were indications of negative additive genetic correlations between DAH and all SBI traits, but because of low prevalence and low heritability of DAH, these results require further scrutiny. Positive additive genetic correlations of rg = 0.37–0.52 were estimated between OFF and withers height and between OFH and withers height, indicating that selection for taller horses will increase disposition to develop OFF and OFH. Selection of broodmares with regards to functional conformation will assist, but cannot replace possible selection against radiographic findings in the limbs of young Warmblood riding horses, particularly with regards to OFF.
doi:10.1186/1297-9686-38-6-657
PMCID: PMC2689269  PMID: 17129565
riding horse; conformation; studbook inspection; radiographic finding; additive genetic correlation

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