G.E.J. Langenbach

Fiber-type composition of the human jaw muscles (Part 1) Origin and functional significance of fiber-type diversity

This is the first of two articles on the fiber-type composition of the human jaw muscles. The present article discusses the origin of fiber-type composition and its consequences. This discussion is presented in the context of the requirements for functional performance and adaptation that are imposed upon the jaw muscles. The human masticatory system must perform a much larger variety of motor tasks than the average limb or trunk motor system. An important advantage of fiber-type diversity, as observed in the jaw muscles, is that it optimizes the required function while minimizing energy use. The capacity for adaptation is reflected by the large variability in fiber-type composition among muscle groups, individual muscles, and muscle regions. Adaptive changes are related, for example, to the amount of daily activation and/or stretch of fibers. Generally, the number of slow, fatigue-resistant fibers is relatively large in muscles and muscle regions that are subjected to considerable activity and/or stretch.

Fiber-type Composition of the Human Jaw Muscles—(Part 2) Role of Hybrid Fibers and Factors Responsible for Inter-individual Variation

This is the second of two articles about fiber-type composition of the human jaw muscles. It reviews the functional relationship of hybrid fibers and the adaptive properties of jaw-muscle fibers. In addition, to explain inter-individual variation in fiber-type composition, we discuss these adaptive properties in relation to environmental stimuli or perturbations. The fiber-type composition of the human jaw muscles is very different from that of limb and trunk muscles. Apart from the presence of the usual type I, IIA, and IIX myosin heavy-chains (MyHC), human jaw-muscle fibers contain MyHCs that are typical for developing or cardiac muscle. In addition, much more frequently than in limb and trunk muscles, jaw-muscle fibers are hybrid, i.e., they contain more than one type of MyHC isoform. Since these fibers have contractile properties that differ from those of pure fibers, this relatively large quantity of hybrid fibers provides a mechanism that produces a very fine gradation of force and movement. The presence of hybrid fibers might also reflect the adaptive capacity of jaw-muscle fibers. The capacity for adaptation also explains the observed large inter-individual variability in fiber-type composition. Besides local influences, like the amount of muscle activation and/or stretch, more general influences, like aging and gender, also play a role in the composition of fiber types.

Dissociation of bone resorption and bone formation in adult mice with a non-functional V-ATPase in osteoclasts leads to increased bone strength

Osteopetrosis caused by defective acid secretion by the osteoclast, is characterized by defective bone resorption, increased osteoclast numbers, while bone formation is normal or increased. In contrast the bones are of poor quality, despite this uncoupling of formation from resorption. To shed light on the effect of uncoupling in adult mice with respect to bone strength, we transplanted irradiated three-month old normal mice with hematopoietic stem cells from control or oc/oc mice, which have defective acid secretion, and followed them for 12 to 28 weeks. Engraftment levels were assessed by flow cytometry of peripheral blood. Serum samples were collected every six weeks for measurement of bone turnover markers. At termination bones were collected for µCT and mechanical testing. An engraftment level of 98% was obtained. From week 6 until termination bone resorption was significantly reduced, while the osteoclast number was increased when comparing oc/oc to controls. Bone formation was elevated at week 6, normalized at week 12, and reduced onwards. µCT and mechanical analyses of femurs and vertebrae showed increased bone volume and bone strength of cortical and trabecular bone. In conclusion, these data show that attenuation of acid secretion in adult mice leads to uncoupling and improves bone strength.

A longitudinal electromyographic study of the postnatal maturation of mastication in the rabbit

At 2 weeks of age, infant rabbits show chewing movements that resemble those of the adult animal. Previous studies have shown that, at that stage, the accompanying masticatory motor pattern is clearly similar to the suckling motor pattern. As early as 4 weeks, chewing muscle activity is indistinguishable from the adult chewing motor pattern. These reports suggest that the adult chewing motor pattern is developed from the suckling motor pattern. In this study, the chewing motor pattern in the intermediate period (between 2 and 4 weeks of age) was investigated by means of fine-wire electromyography and jaw tracking. Maturation of masticatory movements was found to have two phases. Maximum gape increased in the first few days and was followed by strong development of transverse jaw excursions after the age of 17 days. The increase in jaw excursions was brought about by changes in motor behaviour and facilitated by the development of smooth occlusal surfaces. The changes in motor behaviour were: (1) the level of activity of the balancing-side muscles became more equal to that of the working side; (2) the timing of digastric muscle activity became asymmetrical at the age of 17 days; (3) the peak activity of masseter, temporalis, medial pterygoid and lateral pterygoid muscle portions was gradually shifted or prolonged into the power-stroke phase. It can be concluded that the masticatory contraction pattern shifts from one derived from the suckling contraction pattern at the age of 14 days to one almost similar to the adult chewing pattern at the age of 23 days.

Modelling the masticatory biomechanics of a pig

The relationships between muscle tensions, jaw motions, bite and joint forces, and craniofacial morphology are not fully understood. Three‐dimensional (3‐D) computer models are able to combine anatomical and functional data to examine these complex relationships. In this paper we describe the construction of a 3‐D dynamic model using the anatomical (skeletal and muscle form) and the functional (muscle activation patterns) features of an individual pig. It is hypothesized that the model would produce functional jaw movements similar to those recordable in vivo. Anatomical data were obtained by CT scanning (skeletal elements) and MR imaging (muscles). Functional data (muscle activities) of the same animal were obtained during chewing by bipolar intramuscular electrodes in six masticatory muscles and combined with previously published EMG data. The model was driven by the functional data to predict the jaw motions and forces within the masticatory system. The study showed that it is feasible to reconstruct the complex 3‐D gross anatomy of an individuals masticatory system in vivo. Anatomical data derived from the 3‐D reconstructions were in agreement with published standards. The model produced jaw motions, alternating in chewing side, typical for the pig. The amplitude of the jaw excursions and the timing of the different phases within the chewing cycle were also in agreement with previously published data. Condylar motions and forces were within expected ranges. The study indicates that key parameters of the pigs chewing cycle can be simulated by combining general biomechanical principles, individual‐specific data and a dynamic modelling approach frequently used in mechanical engineering.

The adaptive response of jaw muscles to varying functional demands

Jaw muscles are versatile entities that are able to adapt their anatomical characteristics, such as size, cross-sectional area, and fibre properties, to altered functional demands. The dynamic nature of muscle fibres allows them to change their phenotype to optimize the required contractile function while minimizing energy use. Changes in these anatomical parameters are associated with changes in neuromuscular activity as the pattern of muscle activation by the central nervous system plays an important role in the modulation of muscle properties. This review summarizes the adaptive response of jaw muscles to various stimuli or perturbations in the orofacial system and addresses general changes in muscles as they adapt, specific adaptive changes in jaw muscles under various physiologic and pathologic conditions, and their adaptive response to non-surgical and surgical therapeutic interventions. Although the jaw muscles are used concertedly in the masticatory system, their adaptive changes are not always uniform and vary with the nature, intensity, and duration of the stimulus. In general, stretch, increases neuromuscular activity, and resistance training result in hypertrophy, elicits increases in mitochondrial content and cross-sectional area of the fibres, and may change the fibre-type composition of the muscle towards a larger percentage of slow-type fibres. In contrast, changes in the opposite direction occur when neuromuscular activity is reduced, the muscle is immobilized in a shortened position, or paralysed. The broad range of stimuli that affect the properties of jaw muscles might help explain the large variability in the anatomical and physiological characteristics found among individuals, muscles, and muscle portions.

Mineral heterogeneity affects predictions of intratrabecular stress and strain

Knowledge of the influence of mineral variations (i.e., mineral heterogeneity) on biomechanical bone behavior at the trabecular level is limited. The aim of this study is to investigate how this material property affects the intratrabecular distributions of stress and strain in human adult trabecular bone. Two different sets of finite element (FE) models of trabecular samples were constructed; tissue stiffness was either scaled to the local degree of mineralization of bone as measured with microCT (heterogeneous) or tissue stiffness was assumed to be homogeneous. The influence of intratrabecular mineral heterogeneity was analyzed by comparing both models. Interesting effects were seen regarding intratrabecular stress and strain distributions. In the homogeneous model, the highest stresses were found at the surface with a significant decrease towards the core. Higher superficial stresses could indicate a higher predicted fracture risk in the trabeculae. In the heterogeneous model this pattern was different. A significant increase in stress with increasing distance from the trabecular surface was found followed by a significant decrease towards the core. This suggests trabecular bending during a compression. In both models a decrease in strain values from surface to core was predicted, which is consistent with trabecular bending. When mineral heterogeneity was taken into account, the predicted intratrabecular patterns of stress and strain are more consistent with the expected biomechanical behavior as based on mineral variations in trabeculae. Our findings indicate that mineral heterogeneity should not be neglected when performing biomechanical studies on topics such as the (long-term or dose dependent) effects of antiresorptive treatments.

Length changes in the human masseter muscle after jaw movement

The human masseter is a multilayered, complex muscle contributing to jaw motion. Because variations in stretch may cause muscle fibers to function over different portions of their length‐tension curves, the aim of this study was to determine how parts of the masseter lengthen or shorten during voluntary jaw movements made by living subjects. Magnetic resonance (MR) imaging and optically‐based jaw‐tracking were used to measure muscle‐insertion positions for four parts of the muscle with six degrees of freedom (DOF), before and after maximum‐opening, jaw protrusion and laterotrusion in four adult males. Muscle part lengths and intramuscular tendon lengths were calculated, and these data, with fiber‐tendon ratios published previously, were used to estimate putative changes in fiber‐length. During maximum jaw‐opening, the largest increases in muscle length (34–83%) occurred in the medial part of the deep masseter, whereas the smallest changes occurred in the posterior‐most, superficial masseter (2–19%). Smaller changes were found during movement to the ipsilateral side, than during protrusion and movement to the contralateral side. On maximum opening, putative fibers in the deep masseter lengthened up to 83%, whereas those of the superficial masseter stretched up to 72%. The masseter muscle does not stretch uniformly for major jaw movement. Jaw motion to the ipsilateral side causes little length change in any part, and the effect of tendon‐stretch on estimated fiber lengths is not substantial. The stretch that occurs infers there are task‐related changes in the active and passive tensions produced by different muscle regions. Anat Rec 262:293–300, 2001.

Inhibitory regulation of osteoclast bone resorption by signal regulatory protein α

Osteoclasts mediate bone resorption, which is critical for bone development, maintenance, and repair. Proper control of osteoclast development and function is important and deregulation of these processes may lead to bone disease, such as osteoporosis. Previous studies have shown that the cytosolic protein tyrosine phosphatase SHP‐1 acts as a suppressor of osteoclast differentiation and function, but putative inhibitory receptors that mediate recruitment and activation of SHP‐1 in osteoclasts have remained unknown. In the present study, we identify the SHP‐1‐recruiting inhibitory immunoreceptor signal regulatory protein (SIRP) α as a negative regulator of osteoclast activity. SIRPα is expressed by osteoclasts, and osteoclasts from mice lacking the SIRPα cytoplasmic tail and signaling capacity display enhanced bone resorption in vitro. Consequently, SIRPα‐mutant mice have a significantly reduced cortical bone mass. Fur‐thermore, osteoclasts from SIRPα‐mutant mice show an enhanced formation of actin rings, known to be instrumental in bone resorption. SIRPα mutation did not significantly affect osteoclast formation, implying that the role of SIRPα was limited to the regulation of mature osteoclast function. This identifies SIRPα as a bona fide inhibitory receptor that regulates the bone‐resorption activity and supports a concept in which osteoclast function is balanced by the signaling activities of activating and inhibitory immunoreceptors.—Van Beek, E. M., de Vries, T. J., Mulder, L., Schoenmaker, T., Hoeben, K. A., Matozaki, T., Langenbach, G. E. J., Kraal, G., Everts, V., van den Berg, T. K. Inhibitory regulation of osteoclast bone resorption by signal regulatory protein α. FASEB J. 23, 4081‐4090 (2009). www.fasebj.org

Fibre-type composition of rabbit jaw muscles is related to their daily activity.

Skeletal muscles contain a mixture of fibres with different contractile properties, such as maximum force, contraction velocity and fatigability. Muscles adapt to altered functional demands, for example, by changing their fibre‐type composition. This fibre‐type composition can be changed by the frequency, duration and presumably the intensity of activation. The aim of this study was to analyse the relationship between the spontaneous daily muscle activation and fibre‐type composition in rabbit jaw muscles. Using radio‐telemetry combined with electromyography, the daily activity of five jaw muscles was characterized in terms of the total duration of muscle activity (duty time) and the number of activity bursts. Fibre‐type composition of the muscles was classified by analysing the myosin heavy chain content of the fibres. The amount of slow‐type fibres was positively correlated to the duty time and the number of bursts only for activations exceeding 20–30% of the maximum activity per day. Furthermore, cross‐sectional areas of the slow‐type fibres were positively correlated to the duty time for activations exceeding 30% of the maximum activity. The present data indicate that the amount of activation above a threshold (> 30% peak activity) is important for determining the fibre‐type composition and cross‐sectional area of slow‐type fibres of a muscle. Activation above this threshold occurred only around 2% of the time in the jaw muscles, suggesting that contractile properties of muscle fibres are maintained by a relatively small number of powerful contractions per day.