The fine structure of the drum muscles of the Tigerfish,Therapon jarbua,as compared with the trunk musculature

Summary The fibers of drum and trunk muscles of the Tigerfish, Therapon jarbua, differ greatly in diameter. The myofibrils of the trunk muscles are irregularly oriented, while those of the drum muscles are rolled into spiral or concentric bands. Both muscle types possess the sarcomere structure typical of cross-striated musculature. However, the myofibrils of the drum muscles differ greatly in sarcomere length and width from those in the trunk musculature. The trunk muscles contain few mitochondria, whereas in the drum muscles mitochondria are abundant. The sarcoplasmic reticulum (SR) of the drum muscles takes the form of elongated tubes in both the A and the I region; that of the trunk musculature consists of small vesicles. Of the two muscle types, the drum muscle contains more SR. With respect to the form of the T system, the trunk musculature is of the Z type and the drum muscles of the A-I type. The drum muscle displays a considerably greater number of motor endplates; these lack typical junctional folds and have mitochondria with very few cristae. No fat could be demonstrated in either the drum or the trunk muscles. However, the concentration of glycogen is higher in the drum muscle than in the musculature of the trunk.This work was accomplished with support from the Deutsche Forschungsgemeinschaft and is gratefully dedicated to Prof. R. Danneel on the occasion of his 75th birthday.     

A freeze-fracture study of postembryonic differentiation of latissimus dorsi muscles of the chicken

The differentiation of fiber type characteristics in the anterior (ALD) and posterior (PLD) latissimus dorsi muscles is examined by the freeze-fracture technique in 1-, 7- and 30-day-old chicks. Several characteristics of plasma membrane (caveolae, rectilinear arrays, intramembranous particles) and sarcoplasmic reticulum which show fiber type differences in the adult ALD and PLD muscles are compared in the developmental stages. The caveolar density in the ALD fibers is about 20/microns2 at 1 day increasing to about 37/microns2 at 30 days, whereas in the PLD fibers it remains at about 20/microns2 during this period. The distribution of the caveolae in the two muscles is different from the beginning; in the ALD fibers the caveolae are distributed throughout the plasma membrane and in PLD fibers they are patterned into clusters overlying the I band regions. The density of intramembranous particles of 1-day ALD and PLD plasma membranes appears similar, but by 7 days the particle counts in the sarcolemma of the ALD muscle are about twice as numerous as those in the PLD muscle. The rectilinear arrays are virtually absent in the ALD muscle, whereas in the PLD muscle their density is about 10/microns2 at 1 day and about 20/microns2 at 7 days. Already at 1 day posthatching the SR in ALD and PLD fibers has the adult configuration, i.e., an open irregular network in ALD fibers and periodically arranged tubules with triadic expansions in the PLD fibers.(ABSTRACT TRUNCATED AT 250 WORDS)     

Morphological analysis of tendinous structure in the American alligator jaw muscles

In the American alligator, the jaw muscles show seven bundles of tendinous structure: cranial adductor tendon, mandibular adductor tendon, lamina anterior inferior, trap-shaped lamina lateralis, lamina intramandibularis, lamina posterior, and depressor mandibular tendon (originating from the musculus depressor mandibulae, m. pseudotemporalis, m. adductor mandibulae posterior, m. adductor mandibulae externus, m. intramandibularis, m. pterygoideus anterior, and m. pterygoideus posterior). These tendinous structures are composed of many collagen fibrils and elastic fibers; however, the distributions and sizes of the fibers in these tendinous components differ in comparison with those of other masticatory muscles. The differences of these properties reflect the kinetic forces or the stretch applied to each tendon by the muscle during jaw movements in spite of the simple tendon-muscle junctions. © 1993 Wiley-Liss, Inc.     

The effect of fiber-type heterogeneity on optimized work and power output of hindlimb muscles of the salamander Ambystoma tigrinum

Most vertebrate muscles are composed of a mixture of fiber types. However, studies of muscle mechanics have concentrated on homogeneous bundles of fibers. Hindlimb muscles of the tiger salamander, Ambystoma tigrinum, present an excellent system to explore the consequences of fiber heterogeneity. Isometric twitches and work loops were obtained in vitro from two muscles, the m. iliotibialis pars posterior (heterogeneous, containing types I, IIa and IIb fibers) and the m. iliofibularis (nearly homogeneous for type IIa fibers). Maximal isometric twitch and tetanic stresses in m. iliotibialis posterior were significantly greater than in iliofibularis. Work loops were obtained over a range of frequencies (0.5-3.0 Hz) and strains (2-6% muscle length) that encompassed the observed ranges in vivo. Work per cycle from the homogeneous iliofibularis declined from 1.5-3.0 Hz, while that from the heterogeneous m. iliotibialis posterior increased from 0.5 Hz to 2.5 Hz and declined at 3.0 Hz. Power output from the iliofibularis rose with frequency to at least 3 Hz; power from the iliotibialis posterior rose with frequency to 2.5 Hz and declined thereafter. Mass-specific work per cycle and power output were higher in iliofibularis than iliotibialis posterior over most frequencies and strains tested.     

Suboccipital muscles in the cat neck: Morphometry and histochemistry of the rectus capitis muscle complex

The morphometry, histochemistry, and biomechanical relationships of rectus capitis muscles were examined in adult cats. This family of muscles contained six members on the dorsal, ventral, and lateral aspects of the upper cervical vertebral column. Three dorsal muscles (rectus capitis posterior major, medius, and minor) formed a layered complex spanning from C1 and C2 to the skull. Rectus capitis posterior major was composed predominantly of fast fibers, but the other two deeper muscles contained progressively higher proportions of slow fibers. One ventral muscle, rectus capitis anterior major, was architecturally complex. It originated from several cervical vertebrae and appeared to be divided into two different heads. In contrast, rectus capitis anterior minor and rectus capitis lateralis were short, parallel-fibered muscles spanning between the skull and C1. The ventral muscles all had nonuniform distributions of muscle-fiber types in which fast fibers predominated. Dorsal and ventral muscle groupings usually had cross-sectional areas of 0.5 cm² or more, reflecting a potential capacity to generate maximal tetanic force in excess of 9 N. Biomechanical analyses suggested that one muscle, rectus capitis lateralis, had its largest moment in lateral flexion, whereas the other muscles had large, posturally dependent moment arms appropriate for actions in flexion-extension. The observation that most rectus muscles have relatively large cross-sectional areas and high fast-fiber proportions suggests that the muscles may have important phasic as well as postural roles during head movement. © 1993 Wiley-Liss, Inc.     

Induced neuroma formation and target muscle perturbation in the giant fiber pathway of the Drosophila temperature-sensitive mutant shibire

Summary The temperature-sensitive mutation shibire (shi) in Drosophila melanogaster is thought to disrupt membrane recycling processes, including endocytotic vesicle pinch-off. This mutation can perturb the development of nerves and muscles of the adult escape response. After exposure to a heat pulse (6 h at 30° C) at 20 h of pupal development, adults have abnormal flight muscles. Wing depressor muscles (DLM) are reduced in number from the normal six to one or two fibers, and are composed of enlarged fibers that appear to represent fiber fusion; large spaces devoid of muscle fibers suggested fiber deletion. The normal five motor axons are present in the peripheral nerve PDMN near the ganglion. However, while some motor axons pass dorsally to the extant fibers, other motor axons lacking end targets pass into an abnormal posterior branch and terminate in a neuroma, i.e., a tangle of axons and glia without muscle target tissue. Hemisynapses are common in axons of the proximal PDMN and within the neuroma, but they are rarely seen in control (no heat pulse) shi or wild-type flies. All surviving muscle fibers are innervated; no muscle tissue exists without innervation. Fibrillar fine structure and neuromuscular synapses appear normal. Fused fibers have dual innervation, suggesting correct and specific matching of target tissue and motor axons. Motor axons lacking target fibers do not innervate erroneous targets but instead terminate in the neuroma. These results suggest developmental constraints and rules, which may contribute to the orderly, stereotyped development in the normal flight system. The nature of the anomalies inducible in the flight motor system in shi flies implies that membrane recycling events at about 20 h of pupal development are critical to the formation of the normal adult nerve-muscle pattern for DLM flight muscles.     

Myogenesis in two polyclad platyhelminths with indirect development,Pseudoceros canadensis and Stylostomum sanjuania

SUMMARY Myogenesis of two representatives of Platyhelminthes, Stylostomum sanjuania and Pseudoceros canadensis, was followed from egg deposition until well‐differentiated free‐swimming larval stages, using F‐actin staining and confocal laserscanning microscopy. Zonulae adhaerentes are the only structures to stain before 50% of development between egg deposition and hatching in S. sanjuania, and before 67% of development in P. canadenis. Subsequently, irregular fibers appear in the embryo, followed by a helicoid muscle close to the apical pole. Three longitudinal muscle pairs form, of which the dorsal pair remains more pronounced than the others. Gradually, new muscles form by branching or from double‐stranded muscle zones adjacent to existing muscles. This results in an elaborate muscular bodywall that consists of a single helicoid muscle as well as multiple circular and longitudinal muscles. Diverse retractor muscles insert at the sphincter muscles around the stomodeum. The overall arrangement and formation mode of the larval musculature appears very similar in both species, although only P. canadensis has a primary circular muscle posterior to the helicoid muscle. Muscle formation in the apical region of the embryo precedes that at the abapical pole and the primary longitudinal muscles form slightly later than the primary circular muscles. Myogenesis and larval myoanatomy appears highly conserved among polyclad flatworms, but differs significantly from that of other trochozoan clades. Our data suggest that the larval muscular ground pattern of polyclad larvae comprises a bodywall consisting of a helicoid muscle, circular and longitudinal muscles, several retractor muscles, and sphincter muscles around the stomodeum.     

Patterns of musculature as taxonomic characters for the Turbellaria Acoela

While turbellarians are generally assumed to have body-wall musculature consisting routinely of longitudinal, circular, and diagonal fibers, members of the Acoela examined by a fluorescence-microscopy technique specific for actin showed more complicated and distinctive arrangements of muscles, giving promise for better delimiting taxa within this taxonomically difficult order. Certain globose or tear-drop-shaped worms such as Convoluta pulchra and species of Pseudaphanostoma, Mecynostomum, and Otocelis, showed a complex pattern in which muscles longitudinal in the anterior half of the body arc diagonally across the posterior half; complex brushes of parenchymal muscles that cross at the level of the statocyst and arc postero-laterally also characterize these groups. The more elongate acoel Paratomella sp. was found to have musculature dominated by strictly longitudinal fibers and with relatively weak circular fibers and few fibers running diagonally to the body axis, yet the elongate mecynostomid Paedomecynostomum bruneum showed a crossing of antero-longitudinal fibers similar to that seen in the more globose Mecynostomum sp. A distinctive looping of muscles around the mouth is seen in P. bruneum and the Anaperidae. Such similarities and differences in pattern of musculature promise to provide easily recognizable characters for taxonomy of the Acoela at levels ranging from species to family. This revised version was published online in July 2006 with corrections to the Cover Date.     

Immunochemical analysis of troponin T isoforms in adult, embryonic, regenerating, and denervated chicken fast skeletal muscles

Using monoclonal antibodies (McAbs) which can distinguish between breast- and leg-type troponin T (TnT), we studied the spatial distribution of TnT isoforms in adult chicken fast skeletal muscles. The breast (pectoralis major) and leg (iliotibialis posterior) muscles were composed predominantly of homogeneous fibers containing breast- and leg-type TnT, respectively. The posterior latissimus dorsi muscle was composed of heterogeneous fibers of at least two types, namely breast and leg types. In developing and regenerating fast muscles, only leg-type TnT was expressed at early stages, and later breast-type TnT appeared either transiently or permanently. This led ultimately to several distinct adult fast muscle breast/leg TnT isoform profiles. Since both types of TnT were synthesized in embryonic and regenerating muscles with nerves intact as well as in regenerating muscles with nerves resected, the switching on of their expression during fast muscle development appears to be independent of nerves. However, its full development ("fine tuning" of the protein isoform distribution within the fast fiber types) and the maintenance of the adult state are presumed to be dependent on the nerves, since, although regenerating fibers in denervated muscles could exhibit the early and then the later embryonic stainabilities, they again returned to the early embryonic state; further, the denervation of adult muscles caused the replacement of TnT isoform from the adult to the early embryonic state.     

Indentations in the terminal cisternae of amphibian and mammalian skeletal muscle fibers

Indentations (hillocks and dimples) in the terminal cisternae of mammalian and amphibian skeletal muscle fibers were studied using freeze-fracture and serial thin-section techniques. The structures were seen in all muscles and had a regular separation from each other and from the T-tubule. Indentations were smaller than fenestrations and formed concavities in, but not macromolecular pores through, the terminal cisternae. The average numbers of indentations in rat muscles (measured along the length of the terminal cisternae, within 150 nm of the triadic junction) varied from 0.9 per micrometer in soleus fibers to 9.6 per micrometer in posterior cricoarytenoid fibers. The average numbers in amphibian sartorius fibers varied from 1.6 to 3.6 per micrometer in muscles from different species. The regular alignment of the indentations along the triad, as well as a close correlation between their numbers and the contractile properties of the muscle, suggest that they function in contractile activation and may represent sites of calcium release from the terminal cisternae.     

Molecular architecture of muscles in an acoel and its evolutionary implications

We have characterized the homologs of an actin, a troponin I, and a tropomyosin gene in the acoel Symsagittifera roscoffensis. These genes are expressed in muscles and most likely coexpressed in at least a subset of them. In addition, and for the first time for Acoela, we have produced a species-specific muscular marker, an antibody against the tropomyosin protein. We have followed tropomyosin gene and protein expression during postembryonic development and during the posterior regeneration of amputated adults, showing that preexisting muscle fibers contribute to the wound closure. The three genes characterized in this study interact in the striated muscles of vertebrates and invertebrates, where troponin I and tropomyosin are key regulators of the contraction of the sarcomere. S. roscoffensis and all other acoels so far described have only smooth muscles, but the molecular architecture of these is the same as that of striated fibers of other bilaterians. Given the proposed basal position of acoels within the Bilateria, we suggest that sarcomeric muscles arose from a smooth muscle type, which had the molecular repertoire of striated musculature already in place. We discuss this model in a broad comparative perspective.     

Demonstration of primary and secondary muscle fiber architecture of the bovine tongue by diffusion tensor magnetic resonance imaging

The myoarchitecture of the tongue is comprised of a complex array of muscle fiber bundles, which form the structural basis for lingual deformations during speech and swallowing. We used magnetic resonance imaging of the water diffusion tensor to display the primary and secondary fiber architectural attributes of the excised bovine tongue. Fiber orientation mapping provides a subdivision of the tongue into its principal intrinsic and extrinsic muscular components. The anterior tongue consists of a central region of orthogonally oriented intrinsic fibers surrounded by an axially oriented muscular sheath. The posterior tongue consists principally of a central region of extrinsic fibers, originating at the inferior surface and projecting in a fan-like manner in the superior, lateral, and posterior directions, and lateral populations of extrinsic fibers directed posterior-inferior and posterior-superior. Analysis of cross-fiber anisotropy indicates a basic contrast of design between the extrinsic and the intrinsic fibers. Whereas the extrinsic muscles exhibit a uniaxial architecture typical of skeletal muscle, the intrinsic core muscles, comprised of the verticalis and the transversus muscles, show strong cross-fiber anisotropy. This pattern is consistent with the theory that the tongue's core functions as a muscular hydrostat in that conjoint contraction of the transverse and vertical fibers enable the tissue to expand at right angles to these fibers. These findings suggest that three-dimensional analysis of diffusion tensor magnetic resonance imaging provides a structural basis for understanding the micromechanics of the mammalian tongue.     

The fine structure of the somatic muscles and their attachment to the cuticle in two species of Pycnogonida

The fine structure of the somatic muscles and their attachment to the cuticle in the pyenogonids Nymphon (Chaetonymphon) macronyx G. O. Sars and Boreonymphon cf. abyssorum (Norman) is described. The muscles possess characteristics which are typical of arthropod slow muscle fibers: relatively long sarcomeres, a mean A-band length of about 6 μm and a ratio of thin to thick contractile filaments of 4:1. The sarcotubular system consists of distinct t-tubules, an irregular SR part and randomly distributed dyads and triads, the muscles are attached to the cuticle by specialized epidermal cells containing microtubules extending from the cuticular to the muscular side. The myoepidermal and epidermal-cuticular junctions are described.     

Sequential maturation of skeletal muscle groups during postnatal ontogenesis of rats]

Experiments were conducted on rats during the early postnatal period; a study was made of the membrane potential (MP) establishment of the fibers of the skeletal muscles of the neck, the anterior and the posterior limbs. At birth the most mature were the muscles of the neck, and the least -- the muscles of the posterior limb. Establishment of the stationary MP level in the muscles of the neck occurred during the first week after birth, in the muscles of the anterior limbs -- by the 10th-12th day, and of the posterior limbs -- by the 15th-20th day. The order of maturation of various groups of the skeletal muscles was associated with the peculiarities of the neuro-trophic influences at various age periods. Muscles of the neck were characterized at all the developmental stages by a rhythmic low-frequency electromyographic activity. In the muscles of the limbs the rhythmic electromyographic activity was transformed into the discharge high-frequency activity by the period of termination of increase of the MP of the muscles.     

家兔胫骨前肌肌纤维型的分布研究

根据家兔胫骨前肌的肌纤维起止、排列和神经支配特征,将该肌分为前、后两个亚体。利用家兔８例１６侧胫骨前肌,按上述两个亚体分别取材,作恒冷箱冰冻横切,肌球蛋白ＡＴＰ酶染色,将肌纤维分为Ⅰ型、ⅡＡ型、ⅡＢ型,检测各亚体的肌纤维型构成比例,肌束内肌纤维的分布特征,并用图象分析仪测量各亚体肌纤维横切面积和直径。结果发现,前、后亚体以Ⅱ型纤维居多,前亚体ⅡＡ型纤维高达３５．４％,后亚体Ⅰ型纤维多达２４．５％,两者的ⅡＢ型纤维均达５０％左右。而左、右侧之间无差异,肌束周边部内Ⅰ型纤维仅占１２．７～１３．３％,ⅡＢ型纤维高达５９．９～６０．０％,说明受肌束膜压迫影响,ⅡＢ型肌纤维血供少,以适应无氧酵解的功能。各亚体的Ⅰ型纤维较细,Ⅱ型纤维较Ⅱ粗,Ａ型与ⅡＢ型二者相似。作者认为,前亚体主要参与快速有力的足背屈运动,后亚体则维持踝关节的稳定,保持足弓的形状和弹性,以便适应该肌的站立、跑动和跳跃的功能。     

Regeneration of the nervous and muscular system after caudal amputation in the polychaete <Emphasis Type="Italic">Alitta virens</Emphasis> (Annelida: Nereididae)

Regenerating segments in polychaetes offer a vivid example of epimorphic recovery of the lost organs and tissues. It is also a promising object for studying positional information and the mechanisms maintaining the body integrity. With the aim to develop a convenient standardized model, we described the dynamics of recovery of the major anatomical structures and created a staging system for the caudal regeneration in Alitta virens. In average the normal organization of the posterior body end is restored within 10 days after amputation (dpa). The whole regenerative process was divided into 5 stages: (1) wound healing (0–1 dpa), (2) blastema formation (1–2 dpa), (3) patterning and growth of the blastema (2–3 dpa), (4) differentiation of the first regenerated segment (3–5 dpa), (5) formation and differentiation of the subsequent 5–6 segments (5–10 dpa). The regeneration is carried out mainly by epimorphosis, although the elements of intercalary growth as well as the morphallactic transformation of the stump have been noted. Terminal structures of the pygidium (muscles of the anal sphincter, pygidial cavity, pygidial ring nerve, pygidial cirri) appear at stages 1–3, and then (from stage 3) the formation of new metameres begins in front of the pygidium. Differentiation of the first newborn segment is associated with the tissue remodeling in the last old segment. Formation of the next segments resembles accelerated postlarval growth. The neural elements of the regenerative bud are developing faster than the surrounding muscles. The neurites extending from the CNS and PNS come to the surface of the wound epithelium at stage 1. Later, nerve fibers from the CNS lengthen and thicken along with the growth of the regenerative bud. Ganglion, parapodial nerves, oblique muscles and coeloms of the first segment are detected at stage 4. Longitudinal muscles regenerate in anterior to posterior progression, being constantly in contact with the corresponding fibers of the old tissues. All other muscles differentiate from blastemal cells in isolation from the old musculature of the stump. Our data promote the further using of the posterior body end regeneration in A. virens as an experimental model for resolving crucial problems of developmental biology.     

The musculature of horsehair worm larvae (<Emphasis Type="Italic">Gordius aquaticus</Emphasis>,<Emphasis Type="Italic"> Paragordius varius</Emphasis>, Nematomorpha): F-actin staining and reconstruction by cLSM and TEM

The musculature of larvae of Gordius aquaticus was investigated by laser-scanning microscopy and compared to transmission electron microscopic data for the larva of Paragordius varius. In the anterior portion of the body, the preseptum, four different muscle groups can be distinguished: (1) 12 anterior parietal muscles in the body wall, (2) six oblique muscles that function as retractors of the introvert, (3) six proboscideal muscles, which function as retractors for the proboscis, and (4) six muscles associated with spines of the outermost of the three rings of spines. The posterior portion of the body, the postseptum, possesses four pairs of longitudinal muscle strands in G. aquaticus, the postseptal parietal muscles, that are located dorsolaterally and ventrolaterally. These are not clearly visible in P. varius, where instead three pairs of parietal muscles are present. Additional small muscles are associated with the terminal spines and with the duct running from the pseudointestine to the body wall. All fibers show a cross-striated pattern although this striation is less obvious at the ends of the fibers.     

Relationship Between Structure and Mechanical Function of the Tissues of the Intervertebral Joint

This paper reviews our current understanding of the relationshipbetween the structures and properties of the tissues of thespine and their mechanical functions. Emphasis is on the humanlumbar spine. Vertebrae consist of a core of cancellous bone(low density) surrounded by a shell of cortical bone (high stiffness);as a result they have high stiffness but low mass. The intervertebraldisc is able to withstand compression because of the swellingpressure exerted by the nucleus pulposus which is constrained,radially, by the annulus fibrosus. Thus the disc acts as a thick-walledpressure vessel. Collagen fibers within the annulus providereinforcement during compression, bending and torsion of thedisc. Collagen fibers also provide tensile reinforcement andprevent tears spreading across ligaments. The ligamenta flavacontain elastic fibers (low stiffness and low strength) withcollagen fibers (high stiffness and high strength). In the unstretchedligamenta flava, the collagen fibers have almost random orientationsbut they become aligned as the ligament is stretched. This structureenables the high extensibility of elastic fibers to be exploitedbut protects them from damage at high strains. The structureof the interspinous ligament suggests that its main functionis to attach the thoracolumbar fascia to the posterior spine.Thus the fascia is maintained in tension when stretched by theabdominal muscles. This and other observations indicate theimportance of muscles for maintaining the stability of the spinalcolumn.     

The Musculature of <Emphasis Type="Italic">Testudinella patina</Emphasis> (Rotifera: Flosculariacea), Revealed with CLSM

The musculature of Testudinella patina was visualized using phalloidin-linked fluorescent dye by confocal laser scanning microscopy. The conspicuous broad retractors appear to be made up of five separate fibers, of which three anchor in the neck region whereas two extend into the corona. Besides the broad retractors, a total of five paired longitudinal retractors are present and all of them extend into the corona. Incomplete circular muscles are found in groups in the neck region and in the medial and posterior parts of the trunk. The foot musculature comprises eight thin ventral foot muscles and six thicker dorsal foot muscles that all extend from the foot basis to the distal part of the foot. At the basis of the foot, each of the dorsal foot muscles anchors on a smaller, S-shaped subterminal foot muscle. The foot musculature furthermore comprises one pair of paraterminal foot muscles that each anchors basally on a subterminal foot muscle, extends into the most proximal part of the foot and attaches on one of the dorsal foot muscles. The visceral musculature is composed of extremely delicate fibers and is restricted to an area around and posterior to the foot opening. The presence of incomplete circular muscles supports that these muscles are a basal trait for Rotifera, whereas the morphology of the broad retractors and foot muscles is much more specialized and may be autapomorphic for Testudinella or alternatively for this genus and its closest relatives. The present results stress that revealing muscles by staining may produce new information from even well-investigated species, and that this information may contribute to a better understanding of functional as well as phylogenetic aspects of rotifer biology.     

Functional morphology of the cephalopod buccal mass: a novel joint type

The arrangement of the musculature and connective tissues of the buccal mass of the coleoid cephalopods Octopus bimaculoides, Sepia officinalis, and Loliguncula brevis was examined using dissection and histology. Serial sections in three mutually perpendicular planes were used to identify the muscles and connective tissues responsible for beak movements and stability and to describe their morphology and fiber trajectories. Four major beak muscles were identified: the anterior, posterior, superior, and lateral mandibular muscles. The anterior, posterior, and superior mandibular muscles connect the upper beak and the lower beak. Although the lateral mandibular muscles originate on the upper beak, they do not connect to the lower beak and instead insert on a connective tissue sheath surrounding the buccal mass. Examination of the fibers of the lateral mandibular muscles reveals that they have the organization of a muscular hydrostat, with muscle fibers oriented in three mutually perpendicular orientations. Although the beaks are capable of complex opening, closing, and shearing movements, they do not contact one another and are instead connected only by the musculature of the buccal mass. Based on the morphological analysis and observations of freshly dissected beaks undergoing the stereotyped bite cycle, the functional role of the beak muscles is hypothesized. The anterior and superior mandibular muscles are likely responsible for beak closing and shearing movements. The posterior mandibular muscle is likely also involved in beak closing, but may act synergistically with the lateral mandibular muscles to open the beaks. The lateral mandibular muscles may use a muscular-hydrostatic mechanism to control the location of the pivot between the beaks and to generate the force required for beak opening. The lack of contact between the beaks and the morphology of the lateral mandibular muscles suggests that the buccal mass of coleoid cephalopods may represent a previously unexamined flexible joint mechanism. The term "muscle articulation" is proposed here to denote the importance of the musculature in the function of such a joint.