Susan W. Herring

Strain in the Braincase and Its Sutures During Function

The skull is distinguished from other parts of the skeleton by its composite construction. The sutures between bony elements provide for interstitial growth of the cranium, but at the same time they alter the transmission of stress and strain through the skull. Strain gages were bonded to the frontal and parietal bones of miniature pigs and across the interfrontal, interparietal and coronal sutures. Strains were recorded 1) during natural mastication in conjunction with electromyographic activity from the jaw muscles and 2) during stimulation of various cranial muscles in anesthetized animals. Vault sutures exhibited vastly higher strains than did the adjoining bones. Further, bone strain primarily reflected torsion of the braincase set up by asymmetrical muscle contraction; the tensile axis alternated between +45 degrees and -45 degrees depending on which diagonal masseter/temporalis pair was most active. However, suture strains were not related to overall torsion but instead were responses to local muscle actions. Only the coronal suture showed significant strain (tension) during jaw opening; this was caused by the contraction of neck muscles. All sutures showed strain during jaw closing, but polarity depended on the pattern of muscle usage. For example, masseter contraction tensed the coronal suture and the anterior part of the interfrontal suture, whereas the temporalis caused compression in these locations. Peak tensile strains were larger than peak compressive strains. Histology suggested that the skull is bent at the sutures, with the ectocranial surface tensed and the endocranial surface predominantly compressed. Collectively, these results indicate that skulls with patent sutures should be analyzed as complexes of independent parts rather than solid structures.

Hyaluronic acid hydrogels with controlled degradation properties for oriented bone regeneration

Non-healing fractures can result from trauma, disease, or age-related bone loss. While many treatments focus on restoring bone volume, few try to recapitulate bone organization. However, the native architecture of bone is optimized to provide its necessary mechanical properties. Hyaluronic acid (HA) hydrogel scaffold systems with tunable degradation properties were developed for the controlled delivery of osteoinductive and angiogenic growth factors, thus affecting the quantity and quality of regenerated tissue. HA hydrogels were designed to degrade at fast, intermediate, and slow rates due to hydrolysis and further provided controlled release of cationic proteins due to electrostatic interactions. Scaffolds delivering bone morphogenetic protein-2 (BMP-2) were evaluated in a rat calvarial bone critical size defect model. BMP-2 delivery from the HA hydrogels had a clear osteoinductive effect in vivo and, for all hydrogel types, BMP-2 delivery resulted in significant mineralization compared to control hydrogels. The temporal progression of this effect could be modulated by altering the degradation rate of the scaffold. All three degradation rates tested resulted in similar amounts of mineral formation at the latest (six week) time point examined. Interestingly, however, the fastest and slowest degrading scaffolds seemed to result in more organized bone than the intermediate degrading scaffold, which was designed to degrade in 6-8 weeks to match the healing time. Additionally, healing could be enhanced by co-delivery of vascular endothelial growth factor along with BMP-2.

Osteoprotegerin, a Crucial Regulator of Bone Metabolism, Also Regulates B Cell Development and Function

Osteoprotegerin (OPG) is a CD40-regulated gene in B cells and dendritic cells (DCs). We investigated the role of OPG in the immune system by generating opg−/− mice. Like its role as a regulator of bone metabolism, OPG also influences processes in the immune system, notably in B cell development. Ex vivo, opg−/− pro-B cells have enhanced proliferation to IL-7, and in opg−/− spleen, there is an accumulation of type 1 transitional B cells. Furthermore, opg−/− bone marrow-derived DCs are more effective in stimulating allogeneic T cells than control DCs. When challenged with a T-dependent Ag, opg−/− mice had a compromised ability to sustain an IgG3 Ag-specific response. Thus, in the immune system, OPG regulates B cell maturation and development of efficient Ab responses.

Craniofacial Sutures: Morphology, Growth, and In Vivo Masticatory Strains

The growth and morphology of craniofacial sutures are thought to reflect their functional environment. However, little is known about in vivo sutural mechanics. The present study investigates the strains experienced by the internasal, nasofrontal, and anterior interfrontal sutures during masticatory activity in 4–6‐month‐old miniature swine (Sus scrofa). Measurements of the bony/fibrous arrangements and growth rates of these sutures were then examined in the context of their mechanical environment. Large tensile strains were measured in the interfrontal suture (1,036 μϵ ± 400 SD), whereas the posterior internasal suture was under moderate compression (−440 μϵ ± 238) and the nasofrontal suture experienced large compression (−1,583 μϵ ± 506). Sutural interdigitation was associated with compressive strain. The collagen fibers of the internasal and interfrontal sutures were clearly arranged to resist compression and tension, respectively, whereas those of the nasofrontal suture could not be readily characterized as either compression or tension resisting. The average linear rate of growth over a 1‐week period at the nasofrontal suture (133.8 μm, ± 50.9 S.D) was significantly greater than that of both the internasal and interfrontal sutures (39.2 μm ± 11.4 and 65.5 μm ± 14.0, respectively). Histological observations suggest that the nasofrontal suture contains chondroid tissue, which may explain the unexpected combination of high compressive loading and rapid growth in this suture. J. Morphol. 242:167–179, 1999.

Jaw muscles and the skull in mammals: the biomechanics of mastication

Among non-mammalian vertebrates, rigid skulls with tight sutural junctions are associated with high levels of cranial loading. The rigid skulls of mammals presumably act to resist the stresses of mastication. The pig, Sus scrofa, is a generalized ungulate with a diet rich in resistant foods. This report synthesizes previous work using strain gages bonded to the bones and sutures of the braincase, zygomatic arch, jaw joint, and mandible with new studies on the maxilla. Strains were recorded during unrestrained mastication and/or in anesthetized pigs during muscle stimulation. Bone strains were 100-1000 micro epsilon, except in the braincase, but sutural strains were higher, regardless of region. Strain regimes were specific to different regions, indicating that theoretical treatment of the skull as a unitary structure is probably incorrect. Muscle contraction, especially the masseter, caused strain patterns by four mechanisms: (1) direct loading of muscle attachment areas; (2) a compressive reaction force at the jaw joint; (3) bite force loading on the snout and mandible; and (4) movement causing new points of contact between mandible and cranium. Some expected patterns of loading were not seen. Most notably, strains did not differ for right and left chewing, perhaps because pigs have bilateral occlusion and masseter activity.

THE DYNAMICS OF MASTICATION IN PIGS

Abstract Mastication in miniature pigs was studied via microphone recording and correlated electromyography and cinematography. The results were compared with data from the literature on man and the rhesus monkey (Macaca mulatta). It was concluded that anatomical similarities between pigs and higher primates are due to overall correspondences in masticatory systems adapted for processing a wide variety of foodstuffs. The dental crown patterns allow for both initial puncturing and crushing of the bolus and for transverse gliding of the teeth to grind the food. Lateral deviations of the jaw are produced by muscular couples which include the anterior-pulling masseter of one side and the posterior-pulling temporalis of the opposite side. Specific differences between pigs and higher primates are associated with specialization of the former for processing highly resistant foods. Peculiarities of pig mastication include functional independence of crushing and grinding movements, consecutive bilateral grinding, reversing the direction of chewing with each stroke, and a very rapid rate of mastication.

Mechanical Influences on Suture Development and Patency

In addition to their role in skull growth, sutures are sites of flexibility between the more rigid bones. Depending on the suture, predominant loading during life may be either tensile or compressive. Loads are transmitted across sutures via collagenous fibers and a fluid-rich extracellular matrix and can be quasi-static (growth of neighboring tissues) or intermittent (mastication). The mechanical properties of sutures, while always viscoelastic, are therefore quite different for tensile versus compressive loading. The morphology of individual sutures reflects the nature of local loading, evidently by a process of developmental adaptation. In vivo or ex vivo, sutural cells respond to tensile or cyclic loading by expressing markers of proliferation and differentiation, whereas compressive loading appears to favor osteogenesis. Braincase and facial sutures exhibit similar mechanical behavior and reactions despite their different natural environments.

Modulation of mandibular loading and bite force in mammals during mastication

SUMMARY Modulation of force during mammalian mastication provides insight into force modulation in rhythmic, cyclic behaviors. This study uses in vivo bone strain data from the mandibular corpus to test two hypotheses regarding bite force modulation during rhythmic mastication in mammals: (1) that bite force is modulated by varying the duration of force production, or (2) that bite force is modulated by varying the rate at which force is produced. The data sample consists of rosette strain data from 40 experiments on 11 species of mammals, including six primate genera and four nonprimate species: goats, pigs, horses and alpacas. Bivariate correlation and multiple regression methods are used to assess relationships between maximum (ϵ1) and minimum (ϵ2) principal strain magnitudes and the following variables: loading time and mean loading rate from 5% of peak to peak strain, unloading time and mean unloading rate from peak to 5% of peak strain, chew cycle duration, and chew duty factor. Bivariate correlations reveal that in the majority of experiments strain magnitudes are significantly (P<0.001) correlated with strain loading and unloading rates and not with strain loading and unloading times. In those cases when strain magnitudes are also correlated with loading times, strain magnitudes are more highly correlated with loading rate than loading time. Multiple regression analyses reveal that variation in strain magnitude is best explained by variation in loading rate. Loading time and related temporal variables (such as overall chew cycle time and chew duty factor) do not explain significant amounts of additional variance. Few and only weak correlations were found between strain magnitude and chew cycle time and chew duty factor. These data suggest that bite force modulation during rhythmic mastication in mammals is mainly achieved by modulating the rate at which force is generated within a chew cycle, and less so by varying temporal parameters. Rate modulation rather than time modulation may allow rhythmic mastication to proceed at a relatively constant frequency, simplifying motor control computation.)

Patterns of bone strain in the zygomatic arch.

The transmission of force through the skull is complicated by the irregular form of the bones, the interposed sutures, and the multiplicity of loads from the teeth, muscles, and environment. The in vivo relationship between bone strain and muscle function in the mammalian skull is best investigated empirically.

The Role of Canine Morphology in the Evolutionary Divergence of Pigs and Peccaries

Functional differences in the heads of suids (pigs) and tayassuids (peccaries) are examined. Many such differences, including morphology of the craniomandibular joint, complexity of the cheek teeth, and structure of the masticatory muscles, can be correlated with differences in canine anatomy. The evolution of these features in fossil suoids is briefly reviewed. The difference between suids and tayassuids in canine anatomy is mainly one of orientation. In peccaries, the upper canine points downward and can serve as a weapon, whereas in pigs, the upper canine points laterally and is important in visual display. Sexual dimorphism is greater in suids than in tayassuids and is related to population structure. It is suggested that primitive pigs and peccaries had developed different population structures and that the initial canine modification in suids was a response to social selection pressure.