Muscles advance the teeth in sand dollars and other sea urchins

We demonstrate the action of the dental promoter muscles in advancing the continuously growing teeth of sand dollars and sea urchins. Teeth wear at the occlusal end, while new calcite is added to the opposite end. Dental ligaments rigidly hold teeth during chewing, but soften and reform during advancement. The source of forces that advance the teeth was unknown until our discovery of the dental promoter muscles. The muscles, which underly the tooth, attach centrally to the stereom of the pyramid of the Aristotle''s lantern (jaw) and peripherally to a membrane that covers the distal end of the tooth. The muscles shorten along an axis nearly parallel to the long axis of the tooth. We stimulated contraction by addition of acetylcholine, with increasing concentrations of acetylcholine generating higher forces. Forces exerted by this muscle are appropriate for its size and are 1000 times lower than forces exerted by interpyramidal muscles that generate chewing forces. In sand dollars, a single muscle contraction of the dental promoter muscle can account for half the mean daily advancement of the teeth.     

Three-dimensional dynamical capabilities of the human masticatory muscles

Many habitual human jaw movements are non-symmetrical. Generally, it is observed that when the lower incisors move to one side the contralateral condyle moves forwards onto the articular eminence, whereas the ipsilateral condyle stays in the mandibular fossa, moving slightly to the ipsilateral side. These jaw movements are the result of contractions of active masticatory muscles and guided by the temporomandibular joints, their ligaments and passive elastic properties of the muscles. It is not known whether the movements are primarily dependent on passive guidance, active muscle control or both. Therefore, the objective of this study was to analyse the interplay between these factors during non-symmetrical jaw movements. A six-degrees-of-freedom dynamical biomechanical model of the human masticatory system was used. The movements were not restricted to a priori defined joint axes. Jaw movement simulations were performed by unilateral activity of the muscles. The ligaments or the passive elastic properties of the muscles could be removed during these simulations. Laterodeviations conform to naturally observed ones could be generated by unilateral muscle contractions. The movement of the lower incisors was hardly affected by the absence of passive elastic muscle properties or temporomandibular ligaments. The latter, however, influenced the movement of the condyles. The movements could be understood by analysing the combination of forces and torques with respect to the centre of gravity of the lower jaw. In addition, the loading of the condyles appeared to be an important determinant for the movement. This analysis emphasizes that the movements of the jaw are primarily dependent on the orientation of the contributing muscles with respect to this centre of gravity and not on the temporomandibular ligaments or passive elastic muscle properties.     

Mechanical properties of sea-urchin lantern muscles: a comparative investigation of intact muscle groups in Paracentrotus lividus (Lam.) and Stylocidaris affinis (Phil.) (Echinodermata, Echinoidea)

The jaw apparatus, or lantern, of sea-urchins contains five pairs of retractor and protractor muscles which are responsible for lantern displacement. Using intact retractor or protractor groups, the force-length relations of these muscles were compared in two taxonomically distant species, Paracentrotus lividus and Stylocidaris affinis. The total contractile forces generated by the muscles can be resolved into vertical and horizontal components. It was found that the vertical component of the retractors is maximal at a lantern position which is significantly lower (i.e. more protruded) in Paracentrotus than in Stylocidaris. Total forces generated by the retractors were in both species maximal at or above the lantern `resting positions'. In Paracentrotus alone, the total force-displacement curves tended to be bimodal. It is hypothesized that the retractors of Paracentrotus contain two populations of muscle fibres, one adapted for jaw opening and one for lantern retraction. No significant differences in the properties of the protractors of the two species could be identified. The lantern of Paracentrotus is more mobile than that of Stylocidaris and is able to exploit a wider range of food sources. This investigation has shown that the force-length relations of the lantern muscles match their differing working conditions. Accepted: 3 November 1997     

The Aristotle's lantern of the sea-urchin Stylocidaris affinis (Echinoida,Cidaridae): functional morphology of the musculo-skeletal system

Summary The Aristotle's lantern, or masticatory apparatus, of regular sea-urchins is a complex musculo-skeletal system which is thought to have contributed significantly to the evolutionary success of these animals. This paper gives an account of the antomical relationships and functional morphology of both skeletal and soft tissue components in the lantern and related structures of the sea-urchin Stylocidaris affinis (Cidaridae), and compares these features with their equivalent in the previously described lantern of the sea-urchin Paracentrotus lividus (Echinidae, Camarodonta). There are major differences in the skeletons of these lanterns which involve mostly the arrangement and morphology of elements participating in movement, i.e. joints and articular surfaces, and which highlight the generally heavier and less mobile nature of the lantern in the Cidaridae. There are remarkably few differences, however, in the microstructure of the skeletal stereom. Significant dissimilarities were found in the anatomical arrangement of muscles and ligamentous structures and in their macro- and microstructure. The implications of these morphological features for the functioning of the lantern of the Cidaridae are discussed in the context of an integrated model of lantern biomechanics.     

Control exerted by ligaments

The function of the ligaments as local controllers, independent of the central nervous system, in maintaining the integrity of the joint is demonstrated by modelling the human knee in the sagittal plane, and studying its anterior-posterior motion. In addition to the ligaments, the model includes the characteristic geometry of the joint surface and some muscle groups. The connecting reaction forces at the point of contact between the tibia and the femur are considered to be constraint forces due to three different surface motions--gliding, rolling and combined gliding and rolling. It is demonstrated that the ligamentous structure maintains these holonomic and nonholonomic constraints that describe the joint motion, and that stability of the knee joint is provided mainly by ligaments. Muscular structures further stabilize and contribute to joint movement. Computer simulation of rolling movement of the knee is presented to illustrate the importance of the ligaments for joint integrity and stability.     

A biomechanical model on muscle forces in the transfer of spinal load to the pelvis and legs.

Based on musculoskeletal anatomy of the lower back, abdominal wall, pelvis and upper legs, a biomechanical model has been developed on forces in the load transfer through the pelvis. The aim of this model is to obtain a tool for analyzing the relations between forces in muscles, ligaments and joints in the transfer of gravitational and external load from the upper body via the sacroiliac joints to the legs in normal situations and pathology. The study of the relation between muscle coordination patterns and forces in pelvic structures, in particular the sacroiliac joints, is relevant for a better understanding of the aetiology of low back pain and pelvic pain. The model comprises 94 muscle parts, 6 ligaments and 6 joints. It enables the calculation of forces in pelvic structures in various postures. The calculations are based on a linear/non-linear optimization scheme. To gain a better understanding of the function of individual muscles and ligaments, deviant properties of these structures can be preset. The model is validated by comparing calculations with EMG data from the literature. For agonistic muscles, good agreement is found between model calculations and EMG data. Antagonistic muscle activity is underestimated by the model. Imposed activity of modelled antagonistic muscles has a minor effect on the mutual proportions of agonistic muscle activities. Simulation of asymmetric muscle weakness shows higher activity of especially abdominal muscles.     

Contraction and stiffness changes in collagenous arm ligaments of the stalked crinoid Metacrinus rotundus (Echinodermata)

Shortening and stiffness were measured simultaneously in the aboral ligament of arms of sea lilies. Arm pieces were used from which oral tissues (including muscles) were removed, leaving only collagenous ligaments connecting arm ossicles. Chemical stimulation by means of artificial seawater with an elevated concentration of potassium caused both a bending movement and stiffness changes (either softening or stiffening). The movement lasted for 1.5-10 min, and bent posture was maintained. The observation that contraction was not necessarily associated with softening provided evidence against the hypothesis that the shortening of the aboral ligaments was driven by the elastic components that had been charged by the oral muscles and released their strain energy at the softening of the aboral ligaments. The speed of ligamental shortening was slower by at least one order of magnitude than that of muscles. Acetylcholine (10(-5)-10(-3) M) caused both contraction and softening. We conclude that the aboral ligament shows two mechanical activities based on different mechanisms: one is active contraction and the other is connective tissue catch in which passive mechanical properties show mutability. We suggest that there is neural coordination between the two mechanisms.     

Biomechanical model study of pelvic belt influence on muscle and ligament forces

Many patients with low back and/or pelvic girdle pain feel relief after application of a pelvic belt. External compression might unload painful ligaments and joints, but the exact mechanical effect on pelvic structures, especially in (active) upright position, is still unknown. In the present study, a static three-dimensional (3-D) pelvic model was used to simulate compression at the level of anterior superior iliac spine and the greater trochanter. The model optimised forces in 100 muscles, 8 ligaments and 8 joints in upright trunk, pelvis and upper legs using a criterion of minimising maximum muscle stress. Initially, abdominal muscles, sacrotuberal ligaments and vertical sacroiliac joints (SIJ) shear forces mainly balanced a trunk weight of 500N in upright position. Application of 50N medial compression force at the anterior superior iliac spine (equivalent to 25N belt tension force) deactivated some dorsal hip muscles and reduced the maximum muscle stress by 37%. Increasing the compression up to 100N reduced the vertical SIJ shear force by 10% and increased SIJ compression force with 52%. Shifting the medial compression force of 100N in steps of 10N to the greater trochanter did not change the muscle activation pattern but further increased SIJ compression force by 40% compared to coxal compression. Moreover, the passive ligament forces were distributed over the sacrotuberal, the sacrospinal and the posterior ligaments. The findings support the cause-related designing of new pelvic belts to unload painful pelvic ligaments or muscles in upright posture.     

Effects of geometric individualisation of a human spine model on load sharing: neuro-musculoskeletal simulation reveals significant differences in ligament and muscle contribution

In spine research, two possibilities to generate models exist: generic (population-based) models representing the average human and subject-specific representations of individuals. Despite the increasing interest in subject specificity, individualisation of spine models remains challenging. Neuro-musculoskeletal (NMS) models enable the analysis and prediction of dynamic motions by incorporating active muscles attaching to bones that are connected using articulating joints under the assumption of rigid body dynamics. In this study, we used forward-dynamic simulations to compare a generic NMS multibody model of the thoracolumbar spine including fully articulated vertebrae, detailed musculature, passive ligaments and linear intervertebral disc (IVD) models with an individualised model to assess the contribution of individual biological structures. Individualisation was achieved by integrating skeletal geometry from computed tomography and custom-selected muscle and ligament paths. Both models underwent a gravitational settling process and a forward flexion-to-extension movement. The model-specific load distribution in an equilibrated upright position and local stiffness in the L4/5 functional spinal unit (FSU) is compared. Load sharing between occurring internal forces generated by individual biological structures and their contribution to the FSU stiffness was computed. The main finding of our simulations is an apparent shift in load sharing with individualisation from an equally distributed element contribution of IVD, ligaments and muscles in the generic spine model to a predominant muscle contribution in the individualised model depending on the analysed spine level.

     

Anatomy of the feeding apparatus of the nurse shark,Ginglymostoma cirratum

The anatomy of the feeding apparatus of the nurse shark, Ginglymostoma cirratum, was investigated by gross dissection and computer axial tomography. The labial cartilages, jaws, jaw suspension, muscles, and ligaments of the head are described. Palatoquadrate cartilages articulate with the chondrocranium caudally by short, laterally projecting hyomandibulae and rostrally by ethmoorbital articulations. Short orbital processes of the palatoquadrates are joined to the ethmoid region of the chondrocranium by short, thin ethmopalatine ligaments. In addition, various ligaments, muscles, and the integument contribute to the suspension of the jaws. When the mouth is closed and the palatoquadrate retracted, the palatine process of the palatoquadrate is braced against the ventral surface of the nasal capsule and the ascending process of the palatoquadrate is in contact with the rostrodorsal end of the suborbital shelf. When the mandible is depressed and the palatoquadrate protrudes slightly rostroventrally, the palatoquadrate moves away from the chondrocranium. A dual articulation of the quadratomandibular joint restricts lateral movement between the mandible and the palatoquadrate. The vertically oriented preorbitalis muscle spans the gape and is hypothesized to contribute to the generation of powerful crushing forces for its hard prey. The attachment of the preorbitalis to the prominent labial cartilages is also hypothesized to assist in the retraction of the labial cartilages during jaw closure. Separate levator palatoquadrati and spiracularis muscles, which are longitudinally oriented and attach the chondrocranium to the palatoquadrate, are hypothesized to assist in the retraction of the palatoquadrate during the recovery phase of feeding kinematics. Morphological specializations for suction feeding that contribute to large subambient suction pressures include hypertrophied coracohyoideus and coracobranchiales muscles to depress the hyoid and branchial arches, a small oral aperture with well‐developed labial cartilages that occlude the gape laterally, and small teeth. J. Morphol. 241:33–60, 1999. © 1999 Wiley‐Liss, Inc.     

Evaluation of the influence of muscle deactivation on other muscles and joints during gait motion

When any muscle in the human musculoskeletal system is damaged, other muscles and ligaments tend to compensate for the role of the damaged muscle by exerting extra effort. It is beneficial to clarify how the roles of the damaged muscles are compensated by other parts of the musculoskeletal system from the following points of view: From a clinical point of view, it will be possible to know how the abnormal muscle and joint forces caused by the acute compensations lead to further physical damage to the musculoskeletal system. From the viewpoint of rehabilitation, it will be possible to know how the role of the damaged muscle can be compensated by extra training of the other muscles. A method to evaluate the influence of muscle deactivation on other muscles and joints is proposed in this report. Methodology based on inverse dynamics and static optimization, which is applicable to arbitrary motion was used in this study. The evaluation method was applied to gait motion to obtain matrices representing (1) the dependence of muscle force compensation and (2) the change to bone-on-bone contact forces. These matrices make it possible to evaluate the effects of deactivation of one of the muscles of the musculoskeletal system on the forces exerted by other muscles as well as the change to the bone-on-bone forces when the musculoskeletal system is performing the same motion. Through observation of this matrix, it was found that deactivation of a muscle often results in increment/decrement of force developed by muscles with completely different primary functions and bone-on-bone contact force in different parts of the body. For example, deactivation of the iliopsoas leads to a large reduction in force by the soleus. The results suggest that acute deactivation of a muscle can result in damage to another part of the body. The results also suggest that the whole musculoskeletal system must go through extra retraining in the case of damage to certain muscles.     

The pectoral anatomy of selected Ostariophysi. I. The Characiniformes.

The muscles and bones of the pectoral fin of Serrasalmus nattereri, the piranha, resemble those of generalized, lower teleosts with specializations related to a body shape adapted for high-speed carnivory; the pectoral fins being highly mobile with strong ligaments to the rays. The presence of two occipital nerves appears primitive, while the emergence of the subclavian artery within the branchial cavity, as in Gasteropelecus sternicla, appears specialized. The muscles and bones of the latter fish, a fresh-water flying fish, are specialized for self-propelled, aerial flight in the fusion of the right and left girdles greatly expanded for insertions of complex appendicular (flight) muscles, and in the consolidation of the rays and radials into one functional unit moving vertically in flight through contraction of vertical, massive ventral flight muscles. The bony pectoral anatomy of Electrophorus electricus, the electric eel, is specialized in having a mobile joint between the primary girdle and the cleithrum, the former being suspended vertically from the cleithrum by ligaments. The proximal radials and rays are very numerous and vertically aligned. The cleithrum is shaped to accommodate the extensive sternohyoid and pharyngocleithral muscles. The sheet-like appendicular muscles extend beyond the special joint and control its movement. The deeper muscles do not cross this joint. The arterial system is specialized in lacking a deep brachial artery.     

Infraciliature and myoneme system of Campanella umbellaria (Protozoa, Ciliophora, Peritrichida)

The infraciliature and myoneme system of Campanella umbellaria were revealed using the protargol impregnation technique. The main characteristics of the infraciliature are the peristomial ciliary rows (haplokinety and polykineties), which make four and a half turns around the peristomial disc before plunging into the infundibulum, and the aboral infraciliature, which is made up of the aboral ciliary wreath (trochal band) and the scopula. The myoneme system is composed of: 1) longitudinal fibers, which include 60-84 (mean 72.3) short longitudinal fibers, 40-56 (mean 45.8) medium-length longitudinal fibers, and numerous long longitudinal fibers; and 2) circular fibers, which include 8-12 (mean 9.3) peristomial ring fibers, linking fibers, support fibers, and peristomial disc fibers. The various fibers in C. umbellaria are interconnected to form a single myoneme system that may act as a cell skeleton as well as providing the mechanism by which the zooid contracts and relaxes.     

Mechanical properties of the peristomial membrane of the cidaroid sea-urchin Stylocidaris affinis

The peristomial membrane (PM) is the area of flexible body wall which surrounds the mouth of regular sea-urchins. In cidaroid sea-urchins, like Stylocidaris. affnis (Phil.), it contains an extensive endoskeleton of overlapping plates. This paper describes the basic mechanical properties of the PM of S. affinis and compares them with those of the PM of the previously studied echinacean euechinoid Paracentrotus lividus. When subjected to vertical deformation, the PM of S. affinis generates J-shaped tension-deformation curves with an inflexion point at a deformation corresponding to about 3% of the perignathic girdle diameter. It is considerably stiffer than the P. lividus PM and, unlike the latter, it shows asymmetrical properties, i.e. greater resistance to aboral flexion (retraction) than to oral flexion (protraction). In interambulacral regions the junction between the PM endoskeleton and test of S. affinis takes the form of a wide ligament-filled gap which acts as a compliant hinge. In marked contrast to the echinacean PM, that of S. affinis provides little evidence for the presence of mutable collagenous tissue (MCT). The cidaroid PM is interpreted as a design which maximizes mechanical defence whilst retaining the capacity for some flexibility. A possible correlation between the lantern systems and feeding habits of sea-urchins is discussed.     

The lantern of Aristotle: organization of its coelom and origin of its muscles (Echinodermata,Echinoida)

Summary Comparative ultrastructural analyses of the muscles that work the lantern of Aristotle support the opinion that the muscles in question are myoepithelially organized or derivatives of myoepithelia. They are part of the epithelium of the peripharyngeal cavity (=lantern coelom). The coelom epithelium may become multiplelayered in certain regions and is composed of (1) a layer of muscle cells that vary in number and size, (2) nerve cells and their processes that are interspersed between the muscle layer and (3) monociliated adluminal cells that build a continuous cell lining and completely cover the muscle layer. According to their complexity, the lantern muscles exhibit consecutive stages of myoepithelial variations and may finally simulate subepithelial musculature. The results of this study support the hypothesis of a histological development of subepithelial musculature from simple myoepithelia, although both epithelial and mesenchymal musculature may occur in the Echinodermata. Detailed knowledge of the organization of the lantern's coelom space was a prerequisite for the present study. In contrast to previous examinations the lantern coelom is not a continuous space, but is subdivided into several cavities that are partially completely separated from each other. On the one hand, this subdivision is probably caused by the sophisticated arrangement of the lantern's ossicles and on the other by the septa that give rise to muscles that fulfill different functions. lanter's ossicles and on the other by the septa that give     

A comparative study of the life style of two Jurassic irregular echinoids

The irregular echinoids Plesiechinus ornatus (Buckman) (Pygasteroida) and Galeropygus agariciformis (Forbes) (Cassiduloida) occur together in beds of the murchisonoe Zone, Bajocian, outcropping in the Cheltenham region of Gloucestershire. These species were largely restricted to different lithofacies within the carbonate shelf environment. Both adopted a hidden mode of life but achieved this by different techniques. Plesiechinus had fairly short spines and strongly muscular podia over the whole corona and was able to cover itself with coarse substrate particles. The oral tubercles are bilaterally symmetrical and are radially arranged. The oral spines are thought to have pulled sediment out from beneath the test, excavating a small depression for it. Galeropygus bore a dense covering of very small spines and its tube feet were differentiated into aboral respiratory podia and oral suckered podia. It had a preferred anterior direction of locomotion and is thought to have buried itself completely by excavating and ploughing into the substrate as it moved forward. Plesiechinus fed using only its lantern and postulated peristomial tube feet, whereas Galeropygus was a continugus sediment swallower and used its phyllode tube feet and peristomal lip spines in transferring particles towards the mouth.     

A procedure for estimating the relevant forces in the human knee using a four-bar mechanism

Knee injuries, especially those that affect the cruciate and lateral ligaments, are one of the most serious and frequent pathologies that affect the lower human extremity. Hence, the aim of this study is to develop a dynamic model for the lower extremity capable of estimating forces, forces in the cruciate and collateral ligaments and those normal to the articular cartilage, generated in the knee. The proposed model considers a four-bar mechanism in the knee, a spherical joint in the pelvis and a revolute one in the ankle. The four-bar mechanism is obtained by a synthesis process. The dynamic model includes the inertial properties of the femur, tibia, patella and the foot, the ground reaction force and the most important muscles in the knee. Muscle forces are estimated using an optimisation technique. Results from the application of the model on a real human task are presented.     

Evolution of a novel muscle design in sea urchins (Echinodermata: Echinoidea)

The sea urchin (Echinodermata: Echinoidea) masticatory apparatus, or Aristotle's lantern, is a complex structure composed of numerous hard and soft components. The lantern is powered by various paired and unpaired muscle groups. We describe how one set of these muscles, the lantern protractor muscles, has evolved a specialized morphology. This morphology is characterized by the formation of adaxially-facing lobes perpendicular to the main orientation of the muscle, giving the protractor a frilled aspect in horizontal section. Histological and ultrastructural analyses show that the microstructure of frilled muscles is largely identical to that of conventional, flat muscles. Measurements of muscle dimensions in equally-sized specimens demonstrate that the frilled muscle design, in comparison to that of the flat muscle type, considerably increases muscle volume as well as the muscle's surface directed towards the interradial cavity, a compartment of the peripharyngeal coelom. Scanning electron microscopical observations reveal that the insertions of frilled and flat protractor muscles result in characteristic muscle scars on the stereom, reflecting the shapes of individual muscles. Our comparative study of 49 derived "regular" echinoid species using magnetic resonance imaging (MRI) shows that frilled protractor muscles are found only in taxa belonging to the families Toxopneustidae, Echinometridae, and Strongylocentrotidae. The onset of lobe formation during ontogenesis varies between species of these three families. Because frilled protractor muscles are best observed in situ, the application of a non-invasive imaging technique was crucial for the unequivocal identification of this morphological character on a large scale. Although it is currently possible only to speculate on the functional advantages which the frilled muscle morphology might confer, our study forms the anatomical and evolutionary framework for future analyses of this unusual muscle design among sea urchins.     

Effects of lumbo-pelvic rhythm on trunk muscle forces and disc loads during forward flexion: A combined musculoskeletal and finite element simulation study

Previous in-vivo studies suggest that the ratio of total lumbar rotation over pelvic rotation (lumbo-pelvic rhythm) during trunk sagittal movement is essential to evaluate spinal loads and discriminate between low back pain and asymptomatic population. Similarly, there is also evidence that the lumbo-pelvic rhythm is key for evaluation of realistic muscle and joint reaction forces and moments predicted by various computational musculoskeletal models. This study investigated the effects of three lumbo-pelvic rhythms defined based on in-vivo measurements on the spinal response during moderate forward flexion (60°) using a combined approach of musculoskeletal modeling of the upper body and finite element model of the lumbosacral spine. The muscle forces and joint loads predicted by the musculoskeletal model, together with the gravitational forces, were applied to the finite element model to compute the disc force and moment, intradiscal pressure, annular fibers strain, and load-sharing. The results revealed that a rhythm with high pelvic rotation and low lumbar flexion involves more global muscles and increases the role of the disc in resisting spinal loads, while its counterpart, with low pelvic rotation, recruits more local muscles and engages the ligaments to lower the disc loads. On the other hand, a normal rhythm that has balanced pelvic and lumbar rotations yields almost equal disc and ligament load-sharing and results in more balanced synergy between global and local muscles. The lumbo-pelvic rhythm has less effect on the intradiscal pressure and annular fibers strain. This work demonstrated that the spinal response during forward flexion is highly dependent on the lumbo-pelvic rhythm. It is therefore, essential to adapt this parameter instead of using the default values in musculoskeletal models for accurate prediction of muscle forces and joint reaction forces and moments. The findings provided by this work are expected to improve knowledge of spinal response during forward flexion, and are clinically relevant towards low back pain treatment and disc injury prevention.     

Anatomy of the feeding apparatus of the lemon shark,Negaprion brevirostris

The anatomy of the feeding apparatus of the lemon shark, Negaprion brevirostris, is investigated by gross dissection, computer axial tomography, and histological staining. The muscles and ligaments of the head associated with feeding are described. The upper and lower jaws are suspended by the hyoid arch, which in turn is braced against the chondrocranium by a complex series of ligaments. In addition, various muscles and the integument contribute to the suspension and stability of the jaws. The dual jaw joint is comprised of lateral and medial quadratomandibular joints that resist lateral movement of the upper and lower jaws on one another. This is important during feeding involving vigorous head shaking. An elastic ethmoplatine ligament that unites the anterior portion of the upper jaw to the neurocranium is involved with upper jaw retraction. The quadratomandibularis muscle is divided into four divisions with a bipinnate fiber arrangement of the two large superficial divisions. This arrangement would permit a relatively greater force per unit volume and reduce muscle bulging of the jaw adductor muscle in the spatially confined cheek region. Regions of relatively diffuse integumental ligaments overlying the adductor mandibulae complex and the levator palatoquadrati muscle, interspersed with localized regions of longer tendonlike attachments between the skin and the underlying muscle, permit greater musculoskeletal movement relative to the skin. The nomenclature of the hypobranchial muscles is discussed. In this shark they are comprised of the unsegmented coracomandibularis and coracohyoideus, and the segmented coracoarcualis. © 1995 Wiley-Liss, Inc.