Muscle Attachment Sites Research Articles

Morphological Diversity and Evolution of the Jugal in Dinosaurs.

In dinosaurs, as in other reptiles, the homologue of the mammalian zygomatic bone is the jugal. The dinosaurian jugal was primitively triradiate, with posterior, dorsal and anterior processes that respectively contacted the quadratojugal, the postorbital, and the maxilla and lacrimal. However, the jugal evolved along different lines in the three major dinosaurian clades. In theropods this cranial element remained relatively conservative in morphology, apart from being reduced to a rod-like structure in most birds and a few non-avians. In sauropodomorphs the jugal eventually became small, plate-like and nearly restricted to the area below the orbit, even being excluded from the ventral margin of the skull in many derived taxa. Among ornithischians the jugal was highly variable, but in many cases became large and/or adorned with ornamental features such as horns, flanges, and rugosities. The jugal does not appear to have been a site of muscle attachment in most non-avian dinosaurs, but represented an important structural element in the akinetic dinosaurian skull. The conspicuous jugal ornaments seen in many ornithischian dinosaurs, like the less striking ones documented in some saurischians, may have played an important role in the social behavior of the species that possessed them. In many cases they have a weapon-like aspect suggesting use in aggressive displays, if not actual combat, adding to the evidence that agonistic behavior was likely widespread among ornithischians in particular. Anat Rec, 300:30-48, 2017. © 2016 Wiley Periodicals, Inc.

Accuracy of femur reconstruction from sparse geometric data using a statistical shape model

Sparse geometric information from limited field-of-view medical images is often used to reconstruct the femur in biomechanical models of the hip and knee. However, the full femur geometry is needed to establish boundary conditions such as muscle attachment sites and joint axes which define the orientation of joint loads. Statistical shape models have been used to estimate the geometry of the full femur from varying amounts of sparse geometric information. However, the effect that different amounts of sparse data have on reconstruction accuracy has not been systematically assessed. In this study, we compared shape model and linear scaling reconstruction of the full femur surface from varying proportions of proximal and distal partial femur geometry in combination with morphometric and landmark data. We quantified reconstruction error in terms of surface-to-surface error as well as deviations in the reconstructed femur’s anatomical coordinate system which is important for biomechanical models. Using a partial proximal femur surface, mean shape model-based reconstruction surface error was 1.8 mm with 0.15° or less anatomic axis error, compared to 19.1 mm and 2.7–5.6° for linear scaling. Similar results were found when using a partial distal surface. However, varying amounts of proximal or distal partial surface data had a negligible effect on reconstruction accuracy. Our results show that given an appropriate set of sparse geometric data, a shape model can reconstruct full femur geometry with far greater accuracy than simple scaling.

Odyssey in the evolution of a paleopathologist

Abstract. A recent report suggesting perceived limitations of and opportunities in the study of paleopathology suggested the importance of incorporation of scientific methodologies. It seems reasonable to also explore how those methodologies are developed and, indeed, how one approaches paleopathology as a science. The development of one such paleopathologist is delineated from his serendipitous observations to application of hypothesis generation and subsequent testing approach developed during basic medical science education. This approach resulted in recognition of how much he thought he knew was actually contrary to the facts. A critical factor was the collaborative approach with specialists in other fields, wherein linguistic confusion was overcome and perspectives refined by point–counterpoint analysis of hypotheses. The limited reliability of tertiary information was clearly exposed through examination of primary sources – original articles rather than what might be referred to as "meta-analyses". It became clear that linguistics was not the only challenge; application of techniques had to be observed and validated. Without validation one might obtain precision (method repeatedly reveals same results) but at the expense of accuracy (assurance that the method actually assesses the question). Paleontological studies are generally limited to examination of organisms and their traces. Archeologically based studies incorporate additional sources of information (e.g., historic), but are no less subject to such semantic and methodological issues. Proof of concept studies provided new windows to recognition not only of disease but to previous anatomical challenges (e.g., localization of direct muscle attachment sites and distribution). Trans-phylogenetic representation of disease falsified speculation that "evolution" would preclude analysis through time. Pathology is an intrinsic component of life and transcends both species and time. Knowledge gained in a given species and time can be applied to similar disease manifestations in other species in modern time. Once speculations were tested and either verified or falsified, paleo-epidemiologic approach allowed identification of patterns of spread and even application of that knowledge to recognition of human migration patterns. Proof of concept studies provided new windows to recognition not only of disease but to previous anatomical challenges (e.g., localization of direct muscle attachment sites and distribution).

A Common Suite of Coagulation Proteins Function in Drosophila Muscle Attachment.

The organization and stability of higher order structures that form in the extracellular matrix (ECM) to mediate the attachment of muscles are poorly understood. We have made the surprising discovery that a subset of clotting factor proteins are also essential for muscle attachment in the model organism Drosophila melanogaster One such coagulation protein, Fondue (Fon), was identified as a novel muscle mutant in a pupal lethal genetic screen. Fon accumulates at muscle attachment sites and removal of this protein results in decreased locomotor behavior and detached larval muscles. A sensitized genetic background assay reveals that fon functions with the known muscle attachment genes Thrombospondin (Tsp) and Tiggrin (Tig). Interestingly, Tig is also a component of the hemolymph clot. We further demonstrate that an additional clotting protein, Larval serum protein 1γ (Lsp1γ), is also required for muscle attachment stability and accumulates where muscles attach to tendons. While the local biomechanical and organizational properties of the ECM vary greatly depending on the tissue microenvironment, we propose that shared extracellular protein-protein interactions influence the strength and elasticity of ECM proteins in both coagulation and muscle attachment.

Lower limb estimation from sparse landmarks using an articulated shape model

Rapid generation of lower limb musculoskeletal models is essential for clinically applicable patient-specific gait modeling. Estimation of muscle and joint contact forces requires accurate representation of bone geometry and pose, as well as their muscle attachment sites, which define muscle moment arms. Motion-capture is a routine part of gait assessment but contains relatively sparse geometric information. Standard methods for creating customized models from motion-capture data scale a reference model without considering natural shape variations. We present an articulated statistical shape model of the left lower limb with embedded anatomical landmarks and muscle attachment regions. This model is used in an automatic workflow, implemented in an easy-to-use software application, that robustly and accurately estimates realistic lower limb bone geometry, pose, and muscle attachment regions from seven commonly used motion-capture landmarks. Estimated bone models were validated on noise-free marker positions to have a lower (p=0.001) surface-to-surface root-mean-squared error of 4.28mm, compared to 5.22mm using standard isotropic scaling. Errors at a variety of anatomical landmarks were also lower (8.6mm versus 10.8mm, p=0.001). We improve upon standard lower limb model scaling methods with shape model-constrained realistic bone geometries, regional muscle attachment sites, and higher accuracy.

Getting a head in hard soils: Convergent skull evolution and divergent allometric patterns explain shape variation in a highly diverse genus of pocket gophers (Thomomys).

BackgroundHigh morphological diversity can occur in closely related animals when selection favors morphologies that are subject to intrinsic biological constraints. A good example is subterranean rodents of the genus Thomomys, one of the most taxonomically and morphologically diverse mammalian genera. Highly procumbent, tooth-digging rodent skull shapes are often geometric consequences of increased body size. Indeed, larger-bodied Thomomys species tend to inhabit harder soils. We used geometric morphometric analyses to investigate the interplay between soil hardness (the main extrinsic selection pressure on fossorial mammals) and allometry (i.e. shape change due to size change; generally considered the main intrinsic factor) on crania and humeri in this fast-evolving mammalian clade.ResultsLarger Thomomys species/subspecies tend to have more procumbent cranial shapes with some exceptions, including a small-bodied species inhabiting hard soils. Counter to earlier suggestions, cranial shape within Thomomys does not follow a genus-wide allometric pattern as even regional subpopulations differ in allometric slopes. In contrast, humeral shape varies less with body size and with soil hardness. Soft-soil taxa have larger humeral muscle attachment sites but retain an orthodont (non-procumbent) cranial morphology. In intermediate soils, two pairs of sister taxa diverge through differential modifications on either the humerus or the cranium. In the hardest soils, both humeral and cranial morphology are derived through large muscle attachment sites and a high degree of procumbency.ConclusionsOur results show that conflict between morphological function and intrinsic allometric patterning can quickly and differentially alter the rodent skeleton, especially the skull. In addition, we found a new case of convergent evolution of incisor procumbency among large-, medium-, and small-sized species inhabiting hard soils. This occurs through different combinations of allometric and non-allometric changes, contributing to shape diversity within the genus. The strong influence of allometry on cranial shape appears to confirm suggestions that developmental change underlies mammalian cranial shape divergences, but this requires confirmation from ontogenetic studies. Our findings illustrate how a variety of intrinsic processes, resulting in species-level convergence, could sustain a genus-level range across a variety of extrinsic environments. This might represent a mechanism for observations of genus-level niche conservation despite species extinctions in mammals.Electronic supplementary materialThe online version of this article (doi:10.1186/s12862-016-0782-1) contains supplementary material, which is available to authorized users.

Sexual dimorphism in craniomandibular morphology of southern sea otters (Enhydra lutris nereis)

The niche divergence hypothesis suggests that if a species exhibits intersexual differences in diet, selection should favor divergence in the feeding apparatus between the sexes. Recent work revealed that male and female southern sea otters ( Enhydra lutris nereis ) utilize different dietary resources in response to increased population density; females exhibit more specialized diets as a function of smaller home ranges, whereas males exhibit larger home ranges, potentially allowing them to expand their dietary breadths by feeding on prey items that are not found in female home ranges. These dietary differences suggest the potential for sexual dimorphism of the feeding apparatus (i.e., the skull). Here, we tested the hypothesis that male and female southern sea otters exhibit differences in craniomandibular traits directly related to biting ability. Univariate and multivariate analyses of 12 craniomandibular traits showed that size is the primary axis of skull variation, whereas only a handful of craniomandibular traits demonstrated significant shape differences between the sexes. Relative postorbital constriction breadth, masseter in-lever length, and cranial height differed significantly between the sexes. These 3 traits can increase the surface area of jaw muscle attachment sites and thus are directly linked to the mechanics of biting ability. Collectively, these morphological differences indicate that niche divergence may be an important mechanism maintaining sexual dimorphism in southern sea otters.

Entheseal changes and pathological lesions in draught reindeer skeletons – Four case studies from present-day Siberia

Draught use and being ridden often result in typical pathological patterns in animal skeletons. Moreover, physical activity patterns may be reflected in bone biomechanical properties and entheseal changes at muscle attachment sites. This paper presents the pathologies and entheseal changes observed in four draught and/or racing reindeer skeletons against information on their life histories and discusses the probability of linking the observed changes to their use. The results of this study are a useful point of comparison to researchers working on reindeer and other species of draught animals. However, our results also emphasize that entheseal changes and many pathologies have multifactorial etiologies and that interpretation of skeletal change patterns is not straightforward, even when there is information on the life history of the animal and its complete skeleton can be examined.

Morphology of muscle attachment sites in the modern human hand does not reflect muscle architecture.

Muscle attachment sites (entheses) on dry bones are regularly used by paleontologists to infer soft tissue anatomy and to reconstruct behaviors of extinct organisms. This method is commonly applied to fossil hominin hand bones to assess their abilities to participate in Paleolithic stone tool behaviors. Little is known, however, about how or even whether muscle anatomy and activity regimes influence the morphologies of their entheses, especially in the hand. Using the opponens muscles from a sample of modern humans, we tested the hypothesis that aspects of hand muscle architecture that are known to be influenced by behavior correlate with the size and shape of their associated entheses. Results show no consistent relationships between these behaviorally-influenced aspects of muscle architecture and entheseal morphology. Consequently, it is likely premature to infer patterns of behavior, such as stone tool making in fossil hominins, from these same entheses.

IPP Complex Reinforces Adhesion by Relaying Tension-Dependent Signals to Inhibit Integrin Turnover.

Cytoskeleton-mediated forces regulate the assembly and function of integrin adhesions; however, the underlying mechanisms remain unclear. The tripartite IPP complex, comprising ILK, Parvin, and PINCH, mediates the integrin-actin link at Drosophila embryo muscle attachment sites (MASs). Here, we demonstrate a developmentally earlier function for the IPP complex: to reinforce integrin-extracellular matrix (ECM) adhesion in response to tension. In IPP-complex mutants, the integrin-ECM linkage at MASs breaks in response to intense muscle contractility. Mechanistically, the IPP complex is required to relay force-elicited signals that decelerate integrin turnover at the plasma membrane so that the integrin immobile fraction is adequate to withstand tension. Epistasis analysis shows that alleviation of muscle contractility, downregulation of endocytosis, and enhanced integrin binding to the ECM are sufficient to restore integrin-ECM adhesion and maintain integrin-adhesome organization in IPP-complex mutants. Our findings reveal a role for the IPP complex as an essential mechanosensitive regulatory switch of integrin turnover in vivo.

Structure, function and cell dynamics during chaetogenesis of abdominal uncini in Sabellaria alveolata (Sabellariidae, Annelida).

BackgroundDynamic apical microvilli of a single cell, called the chaetoblast, inside an ectodermal invagination form the template of annelid chaetae. Changes in the pattern of microvilli are frozen in time by release of chitin, such that the structure of the definitive chaeta reflects its formation. Cellular interactions during chaetogenesis also influence the structure of the chaeta. Analysing chaetogenesis allows for testing hypotheses on the homology of certain chaetal types. We used this approach to test whether the unusual uncini in Sabellaria alveolata are homologous to apparently similar uncini in other annelid taxa.ResultsOur study reveals unexpected details of sabellariid uncini, which mechanically reinforce the neuropodia enabling their use as paddles. The final structure of the chaeta is caused by pulses of microvilli formation and dynamic interaction between the chaetoblast and adjoining follicle cells. Cell dynamics during chaetogenesis of the uncini in Sabellaria alveolata exceeds by far that reported in previous studies on the formation of this type of chaetae.ConclusionDespite the superficial similarity of uncini in sabellariids and other annelids, differences in structure and details of formation do not support the homology of this type of chaetae. Chaetogenesis of sabellariid uncini involves unexpected microvilli and cell dynamics, and provides evidence that interactions between cells play a larger role in chaetogenesis than previously expected. In addition to their function as anchors, uncini in Sabellaridae stabilize the paddle-shaped notopodia, as each uncinus possesses a long, thin rod that extends deeply into the notopodium. The rods of all uncini in a single row form a bundle inside the notopodium that additionally serves as a muscle attachment site and thus have a similar function to the inner chaeta (acicula) of errant polychaetes (Aciculata).

Global deletion of Panx3 produces multiple phenotypic effects in mouse humeri and femora.

Pannexins form single-membrane channels that allow passage of small molecules between the intracellular and extracellular compartments. Of the three pannexin family members, Pannexin3 (Panx3) is the least studied but is highly expressed in skeletal tissues and is thought to play a role in the regulation of chondrocyte and osteoblast proliferation and differentiation. The purpose of our study is to closely examine the invivo effects of Panx3 ablation on long bone morphology using micro-computed tomography. Using Panx3 knockout (KO) and wildtype (WT) adult mice, we measured and compared aspects of phenotypic shape, bone mineral density (BMD), cross-sectional geometric properties of right femora and humeri, and lean mass. We found that KO mice have absolutely and relatively shorter diaphyseal shafts compared with WT mice, and relatively larger areas of muscle attachment sites. No differences in BMD or lean mass were found between WT and KO mice. Interestingly, KO mice had more robust femora and humeri compared with WT mice when assessed in cross-section at the midshaft. Our results clearly show that Panx3 ablation produces phenotypic effects in mouse femora and humeri, and support the premise that Panx3 has a role in regulating long bone growth and development.

Somatic musculature of rotifers Asplanchna girodi Guerne, 1888 and Trichotria pocillum (Müller, 1776) (Rotifera, Pseudotrrocha, Ploima): Comparative aspect

Comparative study of somatic musculature in illoricate rotifer Asplanchina girodi Guerne, 1888 and loricate Trichotria pocillum (Muller, 1776) has been carried out by the method of phalloidin fluorescence and confocal laser scanning microscopy. Similar layering of muscles is revealed, while significant differences are observed in other aspects. Postcoronal transverse muscle of A. girodi and the dorsal portion of the anterior circular muscle of T. pocillum serve as attachment sites for the refractor muscles. All retractors are formed by smooth muscles or striated muscles, except the lateral retractors of A. girodi, which are formed by the most powerful oblique muscles. In A. girodi there are three pairs of retractors, ten longitudinal muscles, and five circular muscles, with a thick muscular plexus connecting them. In T. pocillum there are four pairs of retractors, five transverse muscles, strong foot retractors, and an arched structure of the head region (new for rotifers). Eight pairs of dorsoventral muscles, as identified in T. pocillum, are completely absent in A. girodi.

Epidermal Cells Expressing Putative Cell Markers in Nonglabrous Skin Existing in Direct Proximity with the Distal End of the Arrector Pili Muscle.

Inconsistent with the view that epidermal stem cells reside randomly spread along the basal layer of the epidermal rete ridges, we found that epidermal cells expressing stem cell markers in nonglabrous skin exist in direct connection with the distal end of the arrector pili muscle. The epidermal cells that express stem cell markers consist of a subpopulation of basal keratinocytes located in a niche at the lowermost portion of the rete ridges at the distal arrector pili muscle attachment site. Keratinocytes in the epidermal stem cell niche express K15, MCSP, and α6 integrin. α5 integrin marks the distal end of the APM colocalized with basal keratinocytes expressing stem cell markers located in a well-protected and nourished environment at the lowermost point of the epidermis; these cells are hypothesized to participate directly in epidermal renewal and homeostasis and also indirectly in wound healing through communication with the hair follicle bulge epithelial stem cell population through the APM. Our findings, plus a reevaluation of the literature, support the hierarchical model of interfollicular epidermal stem cell units of Fitzpatrick. This new view provides insights into epidermal control and the possible involvement of epidermal stem cells in nonmelanoma skin carcinogenesis.

Musculature of the male genitalia in Rivellia (Diptera: Platystomatidae).

The musculature of male genitalia was studied hitherto only in two species of Tephritidae, one species of Platystomatidae, one species of Pallopteridae, and three species of Ulidiidae of the superfamily Tephritoidea. The split of the hypandrium from one structure into three (the hypandrium and two lateral sclerites) is traced. The hypandrial origin of the lateral sclerites of the hypandrial complex is shown based on the localization of muscle attachment sites. The subepandrial origin of the inner lobes of the surstyli is also confirmed.

A Tendon Cell Specific RNAi Screen Reveals Novel Candidates Essential for Muscle Tendon Interaction

Tendons are fibrous connective tissue which connect muscles to the skeletal elements thus acting as passive transmitters of force during locomotion and provide appropriate body posture. Tendon-derived cues, albeit poorly understood, are necessary for proper muscle guidance and attachment during development. In the present study, we used dorsal longitudinal muscles of Drosophila and their tendon attachment sites to unravel the molecular nature of interactions between muscles and tendons. We performed a genetic screen using RNAi-mediated knockdown in tendon cells to find out molecular players involved in the formation and maintenance of myotendinous junction and found 21 candidates out of 2507 RNAi lines screened. Of these, 19 were novel molecules in context of myotendinous system. Integrin-βPS and Talin, picked as candidates in this screen, are known to play important role in the cell-cell interaction and myotendinous junction formation validating our screen. We have found candidates with enzymatic function, transcription activity, cell adhesion, protein folding and intracellular transport function. Tango1, an ER exit protein involved in collagen secretion was identified as a candidate molecule involved in the formation of myotendinous junction. Tango1 knockdown was found to affect development of muscle attachment sites and formation of myotendinous junction. Tango1 was also found to be involved in secretion of Viking (Collagen type IV) and BM-40 from hemocytes and fat cells.

Lobopodians

Lobopodians

Differences in the effects of age on the development of entheseal changes among historical Japanese populations.

This study aims to determine the impacts of both age and physical activity on entheseal changes at sites of muscle and ligament attachment, and proposes that the influence of age on entheseal changes may depend on specific social systems. Entheseal markers of 193 individuals excavated from archaeological sites in Japan were examined, and archaeological evidence from those sites was used to determine the subsistence activities that those individuals engaged in. Fifteenth-century samples were either fishermen or Agehamashiki salt manufacturers, while samples from between the late 17(th) century and the 19(th) century were peasants, townspeople, or samurai. Nineteen entheses were examined using the scoring method of Hawkey and Merbs (1995) to clarify the degree to which age influenced entheseal severity of a given sample as well as the differences in the degree to which age influenced entheseal changes among samples. Results indicated that age influenced entheseal changes; however, the degree to which age influenced entheseal development was dependent on the characteristic physical activities associated with an individual's class and occupation. Furthermore, in certain groups, aging did not significantly impact entheseal changes. The degree to which age influenced entheseal changes was dependent on characteristic physical activities associated with an individual's class and occupation. This indicated that physical activities and lifestyle, along with age, were major factors that contributed to entheseal changes. Based on these results, it was concluded that entheseal changes could serve as a useful tool in the reconstruction of physical activities of past populations.

Larval muscle attachment site (MAS) patterns are a conserved character among Piophilini flies (Diptera, Piophilidae)

The dorsoventral muscle attachment sites (MAS) patterns are described for six species of the tribe Piophilini (Diptera: Piophilidae): Centrophlebomyia furcata (Fabricius), Liopiophila varipes (Meigen), Piophila casei (Linnaeus), Piophila megastigmata McAlpine, Prochyliza nigrimana (Meigen) and Stearibia nigriceps (Meigen). Comparison between the MAS patterns of Piophilini and previous descriptions for Calliphoridae (Diptera) revealed differences in the muscle equipment between the larvae of both taxa. Among the Piophilini, the MAS patterns were highly conserved and only a genus-specific pattern for Piophila species and a species-specific pattern for C. furcata were found. Nevertheless, these differences in MAS patterns were subtle and some intraspecific variability was observed; hence, the MAS patterns do not appear to be suitable as diagnostic characters allowing for species determination of Piophilini larvae.

Muscle attachment site (MAS) patterns for species determination in five species of Sarcophaga (Diptera: Sarcophagidae)

To further establish species determination using the muscle attachment site (MAS) pattern method, third instar larvae of five forensically important species of Sarcophaga Meigen were investigated: Sarcophaga argyrostoma (Robineau-Desvoidy), Sarcophaga caerulescens Zetterstedt, Sarcophaga melanura Meigen, Sarcophaga albiceps Meigen and Sarcophaga similis Meade. As in the previously investigated Calliphoridae, patterns were found to be species specific. The main feature of the Sarcophaga patterns is the divided central horizontal row of segment four. A genus pattern was established to be used as base for comparison in further species determination.