Forelimb Bones Research Articles

Salient features in the locomotion of proboscideans revealed via the differential scaling of limb long bones

The standard differential scaling of proportions in limb long bones (length against circumference) was applied to a phylogenetically wide sample of the Proboscidea, Elephantidae and the Asian (Elephas maximus) and African (Loxodonta africana) elephants. In order to investigate allometric patterns in proboscideans and terrestrial mammals with parasagittal limb kinematics, the computed slopes between long bone lengths and circumferences (slenderness exponents) were compared with published values for mammals, and studied within a framework of the theoretical models of long bone scaling under gravity and muscle forces. Limb bone allometry in E. maximus and the Elephantidae is congruent with adaptation to bending and/or torsion induced by muscular forces during fast locomotion, as in other mammals, whereas the limb bones in L. africana appear to be adapted for coping with the compressive forces of gravity. Hindlimb bones are therefore more compliant than forelimb bones, and the resultant limb compliance gradient in extinct and extant elephants, contrasting in sign to that of other mammals, is shown to be a new important locomotory constraint preventing elephants from achieving a full-body aerial phase during fast locomotion. Moreover, the limb bone pattern of African elephants, indicating a noncritical bone stress not increasing with increments in body weight, explains why their mean and maximal body masses are usually above those for Asian elephants. Differences in ecology may be responsible for the subtle differences observed in vivo between African and Asian elephants, but they appear to be more pronounced when revealed via mechanical patterns dictated by limb bone allometry.

Differential expression of choline kinase isoforms in skeletal muscle explains the phenotypic variability in the rostrocaudal muscular dystrophy mouse

Choline kinase in mammals is encoded by two genes, Chka and Chkb. Disruption of murine Chka leads to embryonic lethality, whereas a spontaneous genomic deletion in murine Chkb results in neonatal forelimb bone deformity and hindlimb muscular dystrophy. Surprisingly, muscular dystrophy isn't significantly developed in the forelimb. We have investigated the mechanism by which a lack of choline kinase beta, encoded by Chkb, results in minimal muscular dystrophy in forelimbs. We have found that choline kinase beta is the major isoform in hindlimb muscle and contributes more to choline kinase activity, while choline kinase alpha is predominant in forelimb muscle and contributes more to choline kinase activity. Although choline kinase activity is decreased in forelimb muscles of Chkb(-/-) mice, the activity of CTP:phosphocholine cytidylyltransferase is increased, resulting in enhanced phosphatidylcholine biosynthesis. The activity of phosphatidylcholine phospholipase C is up-regulated while the activity of phospholipase A(2) in forelimb muscle is not altered. Regeneration of forelimb muscles of Chkb(-/-) mice is normal when challenged with cardiotoxin. In contrast to hindlimb muscle, mega-mitochondria are not significantly formed in forelimb muscle of Chkb(-/-) mice. We conclude that the relative lack of muscle degeneration in forelimbs of Chkb(-/-) mice is due to abundant choline kinase alpha and the stable homeostasis of phosphatidylcholine.

Development of forelimb bones in indigenous sheep fetuses

The study included detection of the sites of ossification centers and their sequence of appearance in the forelimb bones of indigenous sheep fetuses by using double staining method with younger specimens and radiography or maceration methods with old specimens, as well as, histological study with some ages. The results showed that the primary ossification centers of the forelimb in indigenous sheep fetuses appeared firstly in the diaphyses of radius and ulna, humerus, scapula, metacarpus, phalanges and lastly in the carpal bone at an estimated age of 43, 45, 46, 47, 49 - 56 and 90-118 days old respectively. The results of statistical analysis of the total lengths of scapula, humerus, radius, ulna and metacarpus with the lengths of their ossified parts through the 7th – 15th weeks of fetus age, showed presence of significant differences in the average of these measurements among most of studied weeks. Also there was a significant differences in the average of relative increase in the total length and length of ossified part of diaphysis of studied bones during the 7th week in comparison to the same average in the other studied weeks (8th-15th week) of indigenous sheep fetuses age.

Distal Limb Radiography, Ossification, and Growth Plate Closure in the Juvenile Asian Elephant (Elephas maximus)

Eleven juvenile Asian elephants (Elephas maximus) were evaluated radiographically to determine the relative times of growth plate closure and phalangeal ossification in the bones of the distal forelimb. Specifically, the first, second, and third phalanges of the third digit (D3) were evaluated, as well as the third phalanx of digits 1, 2, 4, and 5. All elephants were healthy at the time of examination. A retrospective evaluation of radiographs from six of the 11 juvenile elephants was also completed to augment the data set. This study reports the methods used to obtain high-quality radiographs of the distal juvenile elephant limb, ossification characteristics of the phalanges, relative times of growth plate closure within the proximal phalanges of D3, and a method for age estimation based on radiographic findings. This study will help clinicians in conducting elephant foot radiography, in evaluating foot radiographs in juvenile elephants, in recognizing normal versus pathologic change, and in estimating juvenile elephant age based on radiographic ossification characteristics and growth plate closure times. Consistent use of the proposed foot radiograph technique is recommended to facilitate foot disease recognition and as part of the annual examination of captive Asian elephants.

Accumulations osseuses en périphérie de terriers de petits carnivores : les stigmates de prédation et de fréquentation

Faunal remains from spoil heaps of two burrows inhabited by small carnivores (fox and badger) are analysed from a taphonomical point of view. This analysis provides characteristics for bone accumulation produced by small carnivores and will be a powerful tool for deciphering site formation about the occupational alternation of small carnivores and humans. Identified species were grouped by size classes. Faunal spectrum is composed by varied species of microfauna (70%), mesofauna (30%). Macromammal remains are under represented (less than 1%) and come from scavenged carcasses. Predators and consumed species are compared on the basis of the skeletal part representation, age classes and recording of predation marks (gnawed and digested bones). Skeletal part representation shows that all taxons exhibit a low-representation of axial skeleton and autopodial bones. Predators show a high representation of hind limb bones and a low representation of the fore limb bones whereas consumed species exhibit a reverse pattern. Mortality curve analysis provides an attritional profile for carnivores and helps for the establishment of the season of occupation of the burrow. Adults largely dominate consumed species. Moreover, predation marks are found in 1% of the carnivores’ bones and from 15% to more than 40% on prey bones. The large-sized prey bones only wear gnawed marks (20%) and anthropic marks (10–20%) whereas microfaunal remains exhibits more digested marks (40%) than gnawing stigmata (5%). Medium-sized animals bones wear both marks and with the same proportions (10–15%).

Brief communication: Locomotor limb preferences in captive chimpanzees (Pan troglodytes): implications for morphological asymmetries in limb bones.

Understanding the evolutionary origins of hemispheric specialization remains a topic of considerable interest in a variety of scientific disciplines. Whether nonhuman primates exhibit population-level limb preferences continues to be a controversial topic. In this study, limb preferences for ascending and descending locomotion were assessed as a means of examining the hypothesis that asymmetries in forelimb bones might be attributed to asymmetries in posture. The results indicated that captive chimpanzees showed a population-level leftward asymmetry in descending locomotion but no group bias for ascending locomotion. The results are consistent with previous behavioral studies in captive chimpanzees as well as studies on skeletal asymmetries of the forelimbs of chimpanzees.

Variability in forelimb bone strains during non-steady locomotor activities in goats

The purpose of this study was to investigate the effects of non-steady locomotor activities on load predictability in two goat forelimb bones and to explore the degree to which bone curvature influences load predictability. We measured in vivo bone strains in the radius and metacarpus of juvenile goats performing a variety of natural behaviors in an outdoor arena and compared these strain magnitudes and loading patterns with those measured during steady-state treadmill locomotion. We sought to test two hypotheses: our first hypothesis expects an increase in the variability of strain magnitude and pattern during outdoor non-steady behavior when compared to treadmill locomotion. Our second hypothesis was that the curved radius experiences higher peak strains but less variability during non-steady activities than the straighter metacarpus. We found that unsteady, outdoor locomotion generally caused more variable strain patterns (consistent with the first hypothesis), but not more variable strain magnitudes, than treadmill locomotion in both bones. During outdoor locomotion, higher magnitude strain events in the radius showed more constrained loading patterns than in the metacarpus (consistent with the second hypothesis). In addition to the radius experiencing significantly greater bending strains compared to the straighter metacarpus, these results support the idea of a trade-off between increased predictability of loading pattern and increased bending-induced strain. Strain magnitudes recorded during both outdoor and treadmill locomotion showed a lognormal frequency distribution, but the outdoor bone strain distributions had a greater range because they included high magnitude loading events that did not occur during steady treadmill locomotion.

Regulatory divergence modifies limb length between mammals

Natural selection acts on variation within populations, resulting in modified organ morphology, physiology, and ultimately the formation of new species. Although variation in orthologous proteins can contribute to these modifications, differences in DNA sequences regulating gene expression may be a primary source of variation. We replaced a limb-specific transcriptional enhancer of the mouse Prx1 locus with the orthologous sequence from a bat. Prx1 expression directed by the bat enhancer results in elevated transcript levels in developing forelimb bones and forelimbs that are significantly longer than controls because of endochondral bone formation alterations. Surprisingly, deletion of the mouse Prx1 limb enhancer results in normal forelimb length and Prx1 expression, revealing regulatory redundancy. These findings suggest that mutations accumulating in pre-existing noncoding regulatory sequences within a population are a source of variation for the evolution of morphological differences between species and that cis-regulatory redundancy may facilitate accumulation of such mutations.

Biomechanics of the Bat Limb Skeleton: Scaling, Material Properties and Mechanics

Background/Aims: Wing skeletons of bats are uniquely specialized for flight, reflecting both evolutionary history and the need to maintain structural integrity while generating aerodynamic forces. Methods: We analyzed the anatomical structure of bat wing skeletons in the context of scaling patterns relative to other mammals, material properties and the mechanical function of the wing bones during flight. Results: Compared with nonvolant mammals, the bones of the bat forelimb are elongated, even after correcting for shared phylogenetic history. Bats have consistently larger-diameter bones in the forelimb than do nonvolant mammals but significantly narrower hindlimb bones. Mineralization in the cortical bone of wings is lower than in the long bones of other adult mammals, with a proximodistal gradient of decreasing mineralization. The distal phalanges have only a small amount of mineralized tissue underlying the articular cartilage. Loads required to elicit a 10% length deflection in the wing bones of Glossophaga soricina varied approximately 50-fold along the wing and flexural stiffness nearly 200-fold. Commensurate with low mineralization and flexural stiffness, bat bones experience extraordinarily high bending strains during flight. Conclusion: Bat limb skeletons share features with other mammals and possess specialized characteristics, mostly related to the mechanical demands of flight.

Analysis of forelimb function in basal ceratopsians

AbstractHere, I present the first study of forelimb function in basal ceratopsians [Correction added after publication 28 July 2007: in the preceding sentence ‘Here, I present the first study of forelimb function in basal ceratopsiansDinosauria Orthischia’ was corrected to ‘Here, I present the first study of forelimb function in basal ceratopsians’]. I examined forelimb bones and casts ofPsittacosaurus neimongoliensis,Psittacosaurus mongoliensis,Leptoceratops gracilisandProtoceratops andrewsi. ForPs. neimongoliensisandL. gracilis, I used manual manipulations of bones and casts to determine the range of motion at available forelimb joints. I then used range of motion and morphology to test the predictions of several hypotheses of forelimb function. Forelimb morphology and range of motion indicate thatPsittacosauruswas an obligate biped and thatLeptoceratopsandProtoceratopswere capable of quadrupedal locomotion. Forelimb mobility was too limited inPsittacosaurusfor the hands to reach the mouth.LeptoceratopsandProtoceratopsare members of an evolutionary radiation in which an extension of the glenoid enabled the forelimbs to sprawl laterally for transverse pivoting, perhaps for display, but quadrupedal locomotion was accomplished with the elbows tucked in. InProtoceratops, the radius crosses over the ulna, causing the palms to face caudally. InLeptoceratops, the radius does not cross over the ulna; the palms face largely medially and the fingers have been reoriented so that flexion produces a caudal, propulsive force, even without caudally facing palms.

A nearly complete skeleton of an early juvenile diplodocid (Dinosauria: Sauropoda) from the Lower Morrison Formation (Late Jurassic) of north central Wyoming and its implications for early ontogeny and pneumaticity in sauropods

A nearly complete skeleton of a juvenile sauropod from the Lower Morrison Formation (Late Jurassic, Kimmeridgian) of the Howe Ranch in Bighorn County, Wyoming is described. The specimen consists of articulated mid-cervical to mid-caudal vertebrae and most appendicular bones, but cranial and mandibular elements are missing. The shoulder height is approximately 67 cm, and the total body length is estimated to be less than 200 cm. Besides the body size, the following morphological features indicate that this specimen is an early juvenile; (1) unfused centra and neural arches in presacral, sacral and first to ninth caudal vertebrae, (2) unfused coracoid and scapula, (3) open coracoid foramen, and (4) relatively smooth articular surfaces on the limb, wrist, and ankle bones. A large scapula, short neck and tail and elongate forelimb bones relative to overall body size demonstrate relative growth. A thin-section of the mid-shaft of a femur shows a lack of annual growth lines, indicating an early juvenile individual possibly younger than a few years old. Pneumatic structures in the vertebral column of the specimen SMA 0009 show that pneumatisation of the postcranial skeleton had already started in this individual, giving new insights in the early ontogenetic development of vertebral pneumaticity in sauropods. The specimen exhibits a number of diplodocid features (e.g., very elongate slender scapular blade with a gradually dorsoventrally expanded distal end, a total of nine dorsal vertebrae, presence of the posterior centroparapophyseal lamina in the posterior dorsal vertebrae). Although a few diplodocid taxa, Diplodocus, cf. Apatosaurus, and cf. Barosaurus, are known from several fossil sites near the Howe Ranch, identification of this specimen, even at a generic level, is difficult due to a large degree of ontogenetic variation.

Forelimb segment length proportions in extant hominoids and Australopithecus afarensis

Forelimb proportions have been used to infer locomotor adaptation in Australopithecus afarensis. However, little is known about proportions among individual forelimb segments in extant or fossil hominoids. The partial A. afarensis skeleton A.L. 438-1 and the more complete skeleton A.L. 288-1 provide the opportunity to assess relative length of the arm, forearm, wrist, and palm. We compare scaling relationships between pairs of forelimb bones of extant hominoids and A. afarensis, and length of individual forelimb elements to a body size surrogate. Hylobatids, and to a lesser extent orangutans, have the longest forelimb bones relative to size, although the carpus varies little among taxa, perhaps due to functional constraints of the wrist. Pan species are unique in having long metacarpals relative to ulnar length, demonstrating that they probably differ from the common chimp-human ancestor, and also that developmental mechanisms can be altered to results in differential growth of individual forelimb segments. A. afarensis has no forelimb bones that are significantly longer than those of humans for its size. It falls within the range of variation seen in modern humans for all comparisons relative to size, but appears to differ from the typical human brachial index due to a slightly shorter humerus and/or slightly longer ulna. It has short metacarpals like humans only among hominoids. Thus, while Pan may have elongated its metacarpus relative to ulnar length, A. afarensis may have reduced the length of its metacarpals and possibly its humerus relative to body size from the primitive condition.

Comparison of forelimb function between Deinonychus and Bambiraptor (Theropoda: Dromaeosauridae)

ABSTRACT Fossils and casts of forelimb bones of the dromaeosaurids Deinonychus antirrhopus and Bambiraptor feinbergi were manually manipulated to determine range of motion and to test functional hypotheses. Shoulder motion in Bambiraptor resembles that found by a previous study on Deinonychus. The humerus can be retracted and elevated to subhorizontal positions and protracted somewhat beyond the vertical. In both taxa, the elbow can be strongly flexed but cannot be fully extended. Supination and pronation cannot occur by movement of the radius, which is immobile relative to the ulna. The palms therefore face medially except during wrist extension, which causes obligatory supination. The fingers of Deinonychus remain spread during flexion. In contrast, torsion of the distal articular surface of metacarpal I and the long axis of phalanx III-3 cause the first and third digits of Bambiraptor to approach each other during flexion, the first known instance of opposable fingers in a dinosaur. The morphology and range of motion in the forelimbs of Deinonychus and Bambiraptor enable two-handed prehension with the wrist flexed, one-handed clutching of objects to the chest, use of the hand as a hook, arm-swinging or -raising displays, and use of the forelimbs to maintain balance. Feathered wings, if present, precluded manual apprehension of objects on the ground, two-handed clutching of objects to the chest, and use of digit II to probe crevices. The forelimbs could not be used to dig. Opposability of the fingers of Bambiraptor enabled one-handed prehension, whereas Deinonychus required both hands to hold objects.

Limb development in pouch young of the brushtail possum (Trichosurus vulpecula) and tammar wallaby (Macropus eugenii)

AbstractWe examined the allometric scaling of limb bones, forelimb and hindlimb musculature, and bone mineral content of the brushtail possum Trichosurus vulpecula and tammar wallaby Macropus eugenii eugenii to determine whether the pattern of development of these parameters in early pouch development fits in with the mechanical requirements necessary for true climbing following birth. In the brushtail possum the rates of increase of the forelimb and hindlimb bone widths with body mass were generally significantly lower than those predicted from the hypothesis of ‘static strain similarity’, that is 0.375, and were not significantly different from isometry. Conversely, the rates of increase of limb bone lengths with body mass were significantly greater than predicted, that is >0.25 in this species. In the wallaby the rates of increase in the widths of forelimb bones with body mass were lower than predicted by the hypothesis of static strain similarity whereas those of the hindlimb bones were higher. Again the rates of increase in forelimb and hindlimb bone lengths were all greater than predicted by the static strain hypothesis. These findings show that the neonates of both species had short wide forelimb bones better able to withstand the strain of body mass. Mineral content was lower in the forelimb bones of the pouch young of smaller body size in both species whereas the mineral content of wallaby hindlimb bones was relatively decreased in larger animals. The length of long bones scaled with positive allometry in relation to the associated mass of muscles in the limbs of both species, indicating that greater muscle mass per unit length in the neonate may compensate for the relatively low leverage of the neonatal limb bones. However, the more positive allometric scaling of the length of the hindlimb bones in relation to muscle mass indicates that the neonatal hindlimb has relatively greater muscle mass per unit length of bone. Given the relatively gracile form of the hindlimb bones, this suggests that neonatal forelimb morphology in these two species may result from the timing of birth in relation to limb development rather than the precocious specialization of the forelimb for early locomotion.

Association between findings on palmarodorsal radiographic images and detection of a fracture in the proximal sesamoid bones of forelimbs obtained from cadavers of racing Thoroughbreds

To determine the distribution for limbs and bones in horses with fractures of the proximal sesamoid bones and relationships with findings on palmarodorsal radiographic images. Proximal sesamoid bones obtained from both forelimbs of cadavers of 328 racing Thoroughbreds. Osteophytes; large vascular channels; and fracture location, orientation, configuration, and margin distinctness were categorized by use of high-detail contact palmarodorsal radiographs. Distributions of findings were determined. Relationships between radiographic findings and fracture characteristics were examined by use of chi2 and logistic regression techniques. Fractures were detected in 136 (41.5%) horses. Biaxial fractures were evident in 109 (80%) horses with a fracture. Osteophytes and large vascular channels were evident in 266 (81%) and 325 (99%) horses, respectively. Medial bones typically had complete transverse or split transverse simple fractures, indistinct fracture margins, > 1 vascular channel that was > 1 mm in width, and osteophytes in abaxial wing and basilar middle or basilar abaxial locations. Lateral bones typically had an oblique fracture and distinct fracture margins. Odds of proximal sesamoid bone fracture were approximately 2 to 5 times higher in bones without radiographic evidence of osteophytes or large vascular channels, respectively. Biaxial fractures of proximal sesamoid bones were common in cadavers of racing Thoroughbreds. Differences between medial and lateral bones for characteristics associated with fracture may relate to differences in fracture pathogeneses for these bones. Osteophytes and vascular channels were common findings; however, fractures were less likely to occur in bones with these features.

A Rostrocaudal Muscular Dystrophy Caused by a Defect in Choline Kinase Beta, the First Enzyme in Phosphatidylcholine Biosynthesis

Muscular dystrophies include a diverse group of genetically heterogeneous disorders that together affect 1 in 2000 births worldwide. The diseases are characterized by progressive muscle weakness and wasting that lead to severe disability and often premature death. Rostrocaudal muscular dystrophy (rmd) is a new recessive mouse mutation that causes a rapidly progressive muscular dystrophy and a neonatal forelimb bone deformity. The rmd mutation is a 1.6-kb intragenic deletion within the choline kinase beta (Chkb) gene, resulting in a complete loss of CHKB protein and enzymatic activity. CHKB is one of two mammalian choline kinase (CHK) enzymes (alpha and beta) that catalyze the phosphorylation of choline to phosphocholine in the biosynthesis of the major membrane phospholipid phosphatidylcholine. While mutant rmd mice show a dramatic decrease of CHK activity in all tissues, the dystrophy is only evident in skeletal muscle tissues in an unusual rostral-to-caudal gradient. Minor membrane disruption similar to dysferlinopathies suggest that membrane fusion defects may underlie this dystrophy, because severe membrane disruptions are not evident as determined by creatine kinase levels, Evans Blue infiltration, and unaltered levels of proteins in the dystrophin-glycoprotein complex. The rmd mutant mouse offers the first demonstration of a defect in a phospholipid biosynthetic enzyme causing muscular dystrophy, representing a unique model for understanding mechanisms of muscle degeneration.

Forelimb bone growth and mineral maturation as potential indices of skeletal maturity in sheep

The aim of this study was to characterise the allometric growth and bone mineral maturation of forelimb bones in sheep throughout the growth phase. Forelimb bones (scapula to proximal phalanx) were measured in 84 Merino sheep from similar genetic stock of approximately 12, 32, 64, 84, 116, and 168 weeks of age, with approximately equal numbers of wethers and ewes in each age cohort (n = 14). Sheep were selected for divergence of size, body weight, and condition, in order that the effects of age and body size could be assessed independently. Bone magnesium was measured in the metacarpal and humerus. Results demonstrate the highly coordinated, allometric nature of linear bone growth within the ovine forelimb, though allometric growth patterns differed from those previously published for bone weights. We propose that estimates of maturity proportion (M) based on relative limb bone length or limb proportions may present significant advantages over weight- or composition-based maturity indices, or qualitative variables such as dental eruption or USDA-type maturity scores. Sex differences in growth gradients were minimal, although the higher variability and greater gender divergence of metacarpal bone length casts doubt on the use of its growth plate (breakjoint) to indicate maturity. Bone magnesium content was found to decrease rapidly during the growth period and may represent a useful independent estimate of physiological maturity.

Effect of nutritional restriction and sire genotype on forelimb bone growth and carcass composition in crossbred lambs

The aim of this study was to assess the effect of low or high whole-of-life nutritional planes on bone growth, maturation, and carcass composition in lambs from sires (n = 9) with high estimated breeding values (EBVs) for post-weaning eye muscle depth (PEMD) or liveweight gain (PWWT), compared with sires of industry average for both traits. Lambs (n = 54) were killed at 8 months of age before measurement of forelimb bones, radiographic scoring, and histological measurement of growth plates, and bone ash mineral analysis. A subset of these (n = 36) had carcass composition serially assessed during growth by CAT-scan. Results reveal that the nutritional restriction imposed in this experiment caused significant restriction of skeletal growth, as reflected by shorter, thinner forelimb bones, altered limb proportions, narrowing (and in some cases permanent closure) of growth plates, and an altered bone mineral profile. CAT-scan analysis showed restriction of bone growth was similar to that of muscle growth. Progeny of high muscling (PEMD) sires showed greater muscle growth, but were possibly more susceptible to some of the skeletal effects of nutritional restriction. Greater sire EBVs for PEMD, PWWT, or fat depth were associated with narrower growth plates, suggestive of slower longitudinal bone growth and shorter adult limb length, although bone mass was not affected according to earlier CAT-scan data. Results also suggest that progeny of high PEMD or PWWT sires are earlier maturing in terms of skeletal (or at least limb) growth, although their bone mineral profile (magnesium content) was more consistent with that of physiologically less mature animals.

Genetic and nutritional regulation of lamb growth and muscle characteristics

Combined actions of nutrition and genetic regulation of the growth rate of lambs as well as the physical, biochemical, and eating quality characteristics of their skeletal muscle were assessed in a major field experiment. Data arising were collated and integrated to consolidate findings made at the farm, animal, tissue, cellular, and gene expression levels. At the farm level, increased nutrient availability significantly increased the growth rate of crossbred lambs and increased the growth advantage resulting from the use of sires with high estimated breeding values (EBV) for growth. In contrast, the extra depth of the M. longissimus thoracis et lumborum (EMD) arising from sires with a higher EBV for this trait was constant irrespective of nutrition. Ewe liveweight and body condition were critical in determining the pre-weaning nutrition and growth of lambs, with the LOW plane of nutrition causing stunting of forelimb bones and changes to the allometric growth coefficients for carcass lean and fat. The EBV of the sire for muscling (PEMD) influenced several non-muscling traits, and interactions with nutrition suggested that on HIGH nutrition, absorbed nutrients were partitioned away from wool and fat accretion and towards protein accretion in lambs having superior muscling genetics. Expression levels of known myogenic factors in muscle, together with a suite of peptides and proteins whose identity and levels were determined by proteomic screening, contributed to improved understanding of the mechanisms underpinning nutritional and genetic regulation of skeletal muscle development. The study revealed the need for caution in the use of indirect markers of growth or composition, with their usefulness being constrained if the localisation of the response to selection is specific to parts of the carcass where the selection had initially been concentrated. As well, the possibility that much of the variation explained by potential physiological markers can be accounted for by non-invasive measures of growth and fatness currently used by Australian farmers must be considered.

Remains of Clidastes Cope, 1868, an unexpected mosasaur in the upper Campanian of NW Germany

AbstractThe mosasaurine Clidastes sp. is recognised from cranial and post-cranial remains collected at four localities in NW Germany. Cranial material was found in pelagic turbiditic marls which crop out near the village of Beckum, while post-cranial skeletal elements were collected from sandy limestones exposed near the villages of Schöppingen, Coesfeld and Billerbeck. In stratigraphic order, the units producing these specimens of Clidastes are the Coesfeld, Baumberge and Beckum formations of late Campanian (Late Cretaceous) age. The cranial material comprises the anterior part of a skull and a single isolated tooth, while post-cranial bones comprise a few isolated vertebrae and a partial skeleton including forelimb bones and an articulated vertebral column. Clidastes is known to date from the western North Sea Basin (England), southern Sweden, as well as from North America (Western Interior Seaway and Gulf Coast).