Electromyography of superficial and deep neck muscles during isometric, voluntary, and reflex contractions

Increasingly complex models of the neck neuromusculature need detailed muscle and kinematic data for proper validation. The goal of this study was to measure the electromyographic activity of superficial and deep neck muscles during tasks involving isometric, voluntary, and reflexively evoked contractions of the neck muscles. Three male subjects (28-41 years) had electromyographic (EMG) fine wires inserted into the left sternocleidomastoid, levator scapulae, trapezius, splenius capitis, semispinalis capitis, semispinalis cervicis, and multifidus muscles. Surface electrodes were placed over the left sternohyoid muscle. Subjects then performed: (i) maximal voluntary contractions (MVCs) in the eight directions (45 deg intervals) from the neutral posture; (ii) 50 N isometric contractions with a slow sweep of the force direction through 720 deg; (iii) voluntary oscillatory head movements in flexion and extension; and (iv) initially relaxed reflex muscle activations to a forward acceleration while seated on a sled. Isometric contractions were performed against an overhead load cell and movement dynamics were measured using six-axis accelerometry on the head and torso. In all three subjects, the two anterior neck muscles had similar preferred activation directions and acted synergistically in both dynamic tasks. With the exception of splenius capitis, the posterior and posterolateral neck muscles also showed consistent activation directions and acted synergistically during the voluntary motions, but not during the sled perturbations. These findings suggest that the common numerical-modeling assumption that all anterior muscles act synergistically as flexors is reasonable, but that the related assumption that all posterior muscles act synergistically as extensors is not. Despite the small number of subjects, the data presented here can be used to inform and validate a neck model at three levels of increasing neuromuscular-kinematic complexity: muscles generating forces with no movement, muscles generating forces and causing movement, and muscles generating forces in response to induced movement. These increasingly complex data sets will allow researchers to incrementally tune their neck models' muscle geometry, physiology, and feedforward/feedback neuromechanics.     

Homologies of the transversospinalis muscles in the anterior presacral region of Sauria (crown Diapsida)

Homologies of muscles of the m. transversospinalis group in the dorsal and cervical regions in Sauria are established based on detailed dissections and published accounts of lepidosaurs, crocodylians, and birds. Attachments and directions of tendons comprising this muscle group are fairly conserved among the saurian clades, enabling rather robust inferences on muscle homologies. The innervation pattern indicates that mm. ascendentes are the most lateral muscles of the m. transversospinalis group in Aves, and are inferred to be homologous with the crocodylian m. tendinoarticularis based on their topological similarities. It is suggested here that the lepidosaurian articulo-parietalis part of m. longissimus cervico-capitis actually belongs to the m. transversospinalis group because its tendons of origin are shared with those of m. semispinalis. The avian m. complexus and the lateral part of the crocodylian m. transversospinalis capitis have origins and insertions similar to this lepidosaurian muscle, and are proposed to be homologous with the latter. In some birds, m. longus colli dorsalis, pars profunda continues directly into the anterior cervical region as m. splenius accessorius, suggesting a serially homologous relationship. Similarly, m. splenius anticus continues anteriorly from m. longus colli dorsalis, pars cranialis, and both of these muscles lie dorsal to m. splenius accessorius. Therefore, the currently used nomenclature that regards m. splenius accessorius as a part of m. longus colli dorsalis, pars cranialis and that regards m. splenius anticus as a part of the former muscle does not accurately reflect the serial homologies among these muscles and may not be justified.     

Function and position determine relative proportions of different fiber types in limb muscles of the lizard Tropidurus psammonastes

Skeletal muscles can be classified as flexors or extensors according to their function, and as dorsal or ventral according to their position. The latter classification evokes their embryological origin from muscle masses initially divided during limb development, and muscles sharing a given position do not necessarily perform the same function. Here, we compare the relative proportions of different fiber types among six limb muscles in the lizard Tropidurus psammonastes. Individual fibers were classified as slow oxidative (SO), fast glycolytic (FG) or fast oxidative-glycolytic (FOG) based on mitochondrial content; muscles were classified according to position and function. Mixed linear models considering one or both effects were compared using likelihood ratio tests. Variation in the proportion of FG and FOG fibers is mainly explained by function (flexor muscles have on average lower proportions of FG and higher proportions of FOG fibers), while variation in SO fibers is better explained by position (they are less abundant in ventral muscles than in those developed from a dorsal muscle mass). Our results clarify the roles of position and function in determining the relative proportions of the various muscle fibers and provide evidence that these factors may differentially affect distinct fiber types.     

Low-level exertions of the neck musculature: a study of research methods.

Musculoskeletal neck discomfort is prevalent in many occupations and has been the focus of much research employing surface electromyography (sEMG). Significant differences in experimental methods among researchers make comparisons across studies difficult. The goal of the current research was to use empirical methods to answer specific methodological questions concerning use of sEMG in evaluation of the neck extensor system. This was accomplished in two studies. In Experiment 1, ultrasound technology was used to: (a) determine accessibility of m. splenius and semispinalis capitis with surface electrodes, (b) identify appropriate electrode locations for these muscles/muscle groups, and (c) illustrate potential benefits of using ultrasound in locating muscles/placing electrodes. Experiment 2 sought to assess effects of posture when normalizing sEMG data. Results from Experiment 1 showed no direct access to semispinalis capitis for surface electrodes; their activity can only be sampled as part of a group of muscles. In most subjects, m. splenius was found to be accessible to surface electrodes. Electrode placement recommendations are provided. Results of Experiment 2 showed significant differences in normalized EMG data between a posture-specific technique and a reference posture technique. Posture-specific normalization is recommended for accurately assessing the relative intensity of contractions of these muscles.     

Catalase-positive microperoxisomes in rat soleus and extensor digitorum longus muscle fiber types

The size, distribution, and content of catalase-reactive microperoxisomes were studied cytochemically in slow-twitch oxidative (SO), fast-twitch oxidative glycolytic (FOG), and fast-twitch glycolytic (FG) fibers of soleus and extensor digitorum longus (EDL) rat muscles. Fiber types were classified on the basis of mitochondrial content and distribution, Z-band widths, and myofibril size and shape. Microperoxisomes were generally located between myofibrils at the I-bands. The absence of crystalloid inclusions prevented positive identification of microperoxisomes in nonreacted and aminotriazole-inhibited muscles. EDL and soleus SO fibers possessed the largest microperoxisomes, whereas FOG and FG fibers of the EDL contained small- to medium-sized microperoxisomes. Comparing either microperoxisome number per muscle fiber area or microperoxisome area per fiber area revealed significant differences between fiber types with this ranking: soleus SO greater than EDL SO greater than EDL FOG greater than EDL FG. The present observations demonstrate that the content of catalase-positive microperoxisomes is greatest in the oxidative muscle fiber types. These cytochemical findings account for the higher catalase activity in homogenates of soleus muscles as compared to that of EDL muscles, because the soleus contains more oxidative fibers than EDL.     

Muscle pain induces task-dependent changes in cervical agonist/antagonist activity.

This study examined the effect of experimental neck muscle pain on the EMG-force relationship of cervical agonist and antagonist muscles. Surface EMG signals were detected from the sternomastoid, splenius capitis, and upper trapezius muscles bilaterally from 14 healthy subjects during cervical flexion and extension contractions of linearly increasing force from 0 to 60% of the maximum voluntary contraction (MVC). Measurements were performed before and after injection of 0.5 ml hypertonic and isotonic saline into either the sternomastoid or splenius capitis in two experimental sessions. EMG average rectified value (ARV) of the sternomastoid, splenius capitis, and upper trapezius muscles and the muscle fiber conduction velocity (CV) of the sternomastoid muscle were estimated at 5% MVC force increments. During cervical flexion with injection of hypertonic saline in sternomastoid, ARV of sternomastoid was lower on the side of pain in the force range 25-60% MVC (P < 0.05) and was associated with a bilateral reduction of splenius capitis and upper trapezius ARV (P < 0.01). During cervical extension, injection of hypertonic saline in splenius capitis resulted in lower estimates of splenius capitis ARV on the painful side from 45 to 60% MVC (P < 0.05), which was associated with a bilateral increase in upper trapezius ARV estimates from 50 to 60% MVC (P < 0.001). However, no significant change was identified for estimates of sternomastoid ARV. Experimentally induced neck muscle pain resulted in task-dependent changes in cervical agonist/antagonist activity without modifications in muscle fiber CV.     

Morphometry of Macaca mulatta forelimb. II. Fiber-type composition in shoulder and elbow muscles

The present study examined the fiber-type proportions of 22 muscles spanning the shoulder and/or elbow joints of three Macaca mulatta. Fibers were classified as one of three types: fast-glycolytic (FG), fast-oxidative-glycolytic (FOG), or slow-oxidative (SO). In most muscles, the FG fibers predominated, but proportions ranged from 25-67% in different muscles. SO fibers were less abundant except in a few deep, small muscles where they comprised as much as 56% of the fibers. Cross-sectional area (CSA) of the three fiber types was measured in six different muscles. FG fibers tended to be the largest, whereas SO fibers were the smallest. While fiber-type size was not always consistent between muscles, the relative size of FG fibers was generally larger than FOG and SO fibers within the same muscle. When fiber CSA was taken into consideration, FG fibers were found to comprise over 50% of the muscle's CSA in almost all muscles.     

Comparative histochemical study of prosimian primate hindlimb muscles. II. Populations of fiber types

The populations of fiber types in hindlimb muscles of the tree shrew (Tupaia glis), lesser bushbaby (Galago senegalensis), and the slow loris (Nycticebus coucang) were described and an attempt was made to correlate populations of fiber types and locomotor patterns. Muscle fibers were assigned to one of the following groups: fast-twitch glycolytic (FG), fast-twitch oxidative-glycolytic (FOG), and slow-twitch oxidase (SO). Histochemical techniques for the demonstration of alkaline- and acid-stable ATPase, succinic dehydrogenase, and mitochondrial alpha-glycerophosphate dehydrogenase were used in the classification of muscle fibers. Results indicated that the FG fiber type is the predominant fiber type in muscles used for jumping, the FOG fiber type is predominant in muscles used for running, and the SO fiber type occurs in high percentages in postural muscles. The SO fiber was also the most common fiber in muscles of the slow loris-a species that exhibits a slow, deliberate, sustained locomotor pattern. Intramuscular regional variations in populations were seen in some larger muscles of the tree shrew, but not in the lesser bushbaby and slow loris. Our results did not support the contentions of others that analogous muscles in different species have similar populations of fiber types.     

Comparative histochemical study of prosimian primate hindlimb muscles. I. Muscle fiber types

The profiles of fiber types in hindlimb muscles from the tree shrew (Tupaia glis), lesser bushbaby (Galago senegalensis), and the slow loris (Nycticebus coucang) were determined using histochemical techniques. Fibers were classified as fast-twitch oxidative-glycolytic (FOG), fast-twitch glycolytic (FG), slow-twitch oxidative (SO), or fast-twitch oxidative (FO), according to reactions for alkaline-stable ATPase, acid-stable ATPase, alpha-glucan phosphorylase, reduced nicotinamide adenine dinucleotide diaphorase, succinate dehydrogenase, mitochondrial alpha-glycerophosphate dehydrogenase (MaGPDH), and beta-hydroxybutyric dehydrogenase, as well as glycogen staining by the periodic acid-Schiff technique. Prolonged dissection of numerous muscles was carried out on hindlimbs submersed in cold Tyrode's solution; such treatment had no qualitative effect on enzyme staining reactions, but it is not a suitable procedure if one wishes to stain for glycogen. Fast-twitch oxidative (FO) fibers are alkaline-stable ATPase-positive and possess low MalphaGPDH enzyme activity. These fibers have not been reported previously in any hindlimb muscles. No muscles of any species studies were homogeneous with respect to fiber type. Slow loris muscles lacked FG fibers. The majority of the muscles of the slow loris contained numerous SO fibers. The relationship between enzyme activities and locomotor pattern is discussed.     

Fiber type homogeneity of the flight musculature in small birds

Studies of medium- and large-bodied avian species have suggested that variation in flight muscle composition is related to differences in flight behavior. For example, slow-twitch or tonic fibers are generally found only in the flight muscles of non-volant or soaring/gliding birds. However, we know comparatively little about fiber composition of the muscles of the smallest birds. Here we describe the fiber composition of muscles from the wings, shoulders, and legs of two small avian species, which also display very high wingbeat frequencies: Anna's hummingbirds (Calypte anna) and zebra finches (Taeniopygia guttata). All flight muscles examined in both species contained exclusively fast oxidative glycolytic (FOG) fibers. These unique results suggest that fast oxidative fibers are both necessary and sufficient for the full range of flight behaviors in these small-bodied birds. Like all other studied birds, the zebra finch gastrocnemius, a tarsometatarsal extensor, contained a mixture of FOG (27.1%), slow oxidative (SO, 12.7%), and fast glycolytic (FG, 60.2%) fibers. By contrast, the hummingbird gastrocnemius lacked FG fibers (85.5% FOG, 14.5% SO), which may reflect the reduced role of the hindlimb during take-off. We further hypothesize that thermogenic requirements constrain fiber type heterogeneity in these small endothermic vertebrates.     

Further studies of the clinical anatomy of the craniocervical region

In this publication, some special aspects of the topography in the nuchal region were examined as anatomical support for the dorsal approach to the posterior cranial fossa. The suboccipital muscles rectus capitis posterior minor and rectus capitis lateralis were measured concerning important characteristics as for instance their length and width. The proportions of some muscles arising from the atlas and the relations of the vertebral artery to defined points were evaluated.     

Ultrastructural localization of lectin receptors on the bone-marrow sinusoidal endothelium of the rat

The purpose of this study was to estimate the absolute and relative masses of the three types of skeletal muscle fibers in the total hindlimb of the male Sprague-Dawley rat (Rattus norvegicus). For six rats, total body mass was recorded and the following weights taken from dissection of one hindlimb: 32 individual major muscles or muscle parts, remaining skeletal musculature (small hip muscles and intrinsic foot muscles), bone, inguinal fat pad, and skin. The fibers from the 32 muscles or muscle parts (which constituted 98% of the hindlimb skeletal muscle mass) were classified from histochemistry as fast-twitch oxidative glycolytic (FOG), fast-twitch glycolytic (FG), or slow-twitch oxidative (SO), and their populations were determined. Fiber cross-sectional areas from the same muscles were measured with a digitizer. Mass of each of the fiber types within muscles and in the total hindlimb was then calculated from fiber-type population, fiber-type area, and muscle-mass data. Skeletal muscle made up 71% of the total hindlimb mass. Of this, 76% was occupied by FG fibers, 19% by FOG fibers, and 5% by SO fibers. Thus, the FG fiber type is clearly the predominant fiber type in the rat hindlimb in terms of muscle mass. Fiber-type mass data are compared with physiological (blood flow) and biochemical (succinate dehydrogenase activities) data for the muscles taken from previous studies, and it is demonstrated that these functional properties are closely related to the proportions of muscle mass composed of the various fiber types.     

Fiber subtype distribution of pharyngeal dilator muscles and diaphragm in the cat

In previous studies differences were frequently found between the pharyngeal dilator muscles and the thoracic respiratory muscles in their patterns of electrical and mechanical activity during the respiratory cycle, with both resting and stimulated breathing. However, little is known about the intrinsic properties of the pharyngeal muscles and how they relate to the intrinsic properties of the diaphragm. In the present study, the fiber subtype distributions of two pharyngeal dilator muscles, the geniohyoid and the sternohyoid, were ascertained histochemically in the cat. The geniohyoid and the sternohyoid muscles had a preponderance of fast glycolytic (FG) fibers (mean 48 and 55%, respectively), a smaller number of fast oxidative-glycolytic (FOG) fibers (mean 36 and 31%, respectively), and few slow oxidative (SO) fibers (mean 16 and 14%, respectively). The percentages of SO fibers of both hyoid muscles were significantly (P less than 0.01) lower than that of the costal diaphragm, and the percentages of FOG and FG fibers were significantly higher than that of the diaphragm. In conclusion, the geniohyoid and sternohyoid muscles have histochemical characteristics usually associated with fast contraction and intermediate endurance properties.     

Motor units of the primary ankle extensor muscles of the opossum (Didelphis virginiana): functional properties and fiber types

Motor units of the medial gastrocnemius (MG) and the single lateral gastrocnemius/soleus (LG/S) muscles of the opossum (Didelphis virginiana) were found to have uniformly slow contraction times relative to homologous muscles of the cat. Though a broad range of peak tetanic tensions was found among motor units from both muscles, most of the motor units were quite large relative to tension of the whole muscle. Comparison of the relative sizes of motor units showed that those of LG/S are significantly larger and slower than the units of MG. This suggests that the motor units of the two muscles may be differentially recruited during different behaviors. All of the MG and LG/S motor units were highly or moderately resistant to fatigue. Histochemical staining for NADH-diaphorase activity indicated consistently high levels of the enzyme in all of the fibers of both muscles. Apparently, all of the fast motor units consist of fast oxidative/glycolytic (FOG)-type muscle fibers. Our data provide functional evidence that the types of myofibrillar ATPase demonstrated by Brooke and Kaiser ('70), are not necessarily correlated to physiological classification of fiber types as slow oxidative (SO), fast oxidative/glycolytic (FOG), and fast glycolytic (FG) (Peter et al., '72). Perhaps compartmentalization of muscle fiber types may be a first step in the separation of muscles into multiple heads during the evolution of specialization to diverse locomotor habits among the mammals.     

New observations on the splenius capitis and rectus capitis ventralis muscles of the Common Swift Apus apus (Apodidae)

In swifts and hummingbirds (Apodiformes), the splenius capitis muscle displays a characteristic modification, the 'cruciform origin'. The muscle pairs arise from the second vertebra by several slips which criss-cross and interdigitate with each other, before inserting into the skull. In the course of a study on the neck muscles of the Common Swift, we paid special attention to the arrangement of these slips of the splenius capitis muscle, and noted a previously unrecognized individual modification of this muscle. In addition, we observed an incipient cruciform origin of the deep portion (slip) of the rectus capitis ventralis muscle that has not been noticed for swifts or any other avian taxon before. The development and function of these modifications of the splenius capitis and rectus capitis ventralis muscle are discussed.     

Oxygen uptake,heart rate and activities of locomotor muscles during a critical swimming speed protocol in the gilthead sea bream Sparus aurata

Oxygen uptake, heart rate and contraction frequencies of slow oxidative (SO) and fast glycolytic (FG) muscle were measured simultaneously in gilthead seabream Sparus aurata submitted to stepwise increases in current speed in a swimming respirometer. Variation in oxygen uptake was closely related to variation in heart rate, over initial steps these rose in concert with an increase in contraction frequency of SO muscle. There was an asymptote in oxygen uptake and heart rate at high speeds that reflected a transition from exclusive use of aerobic SO muscle to a combination of SO and anaerobic FG muscle, and which preceded fatigue.     

Structural fiber reinforcement of keel blubber in harbor porpoise (Phocoena phocoena)

This study investigated the functional morphology of the blubber that forms the caudal keels of the harbor porpoise (Phocoena phocoena). Blubber is a pliant biocomposite formed by adipocytes and structural fibers composed of collagen and elastic fibers. Caudal keels are dorsally and ventrally placed triangular wedges of blubber that define the hydrodynamic profile of the porpoise tailstock. Mechanical tests on carcasses demonstrate that when keels are bent, they strain nonuniformly along their lengths, with highest strains just caudal to the dorsal fin and lowest at the insertion of the flukes. Therefore, caudal keels undergo nonuniform longitudinal deformation while maintaining a stable, triangular cross-sectional shape. Polarizing and transmitted light microscopy techniques were used to investigate blubber's 3D fiber architecture along the length of the dorsal keel. The triangular cross-sectional shape of the keel appears to be maintained by structural fibers oriented to act as tensile stays. The construction of the blubber composite is regionally specific :structural fiber densities and diameters are higher in the relatively stiff caudal region of the keel than in the more deformable cranial keel region. The orientations of structural fibers also change along the length of the keel. Cranially, no fibers are oriented along the long axis, whereas a novel population of longitudinally oriented fibers reinforces the keel at the insertion of the flukes. Thus, differences in the distribution and orientation of structural fibers contribute to the regionally specific mechanical properties of the dorsal keel.     

Sexual dimorphism in histological characteristics and contractility of the iliofibularis muscle in the lizard Sceloporus torquatus

The iliofibularis is a hindlimb muscle used in lizard locomotion that is composed of at least three types of fibres: fast‐twitch‐glycolytic (FG), fast‐twitch‐oxidative‐glycolytic fibre (FOG) and slow‐twitch‐oxidative (SO). The striated skeletal muscle is a highly plastic tissue undergoing phenotypic change in response to activity. The lizard Sceloporus torquatus has sexual differences associated with microhabitat use, which can be reflected in the physiology and anatomy of the muscle, and thus, in our study, we analysed the morphological and contractile characteristics of the iliofibularis muscle (IF) of S. torquatus males and females. We found a larger prevalence of FOG compared with FG and SO fibres in the muscle of both sexes. We also found that males show larger areas of the three types of fibres, develop greater strength but also faster fatigue than females, suggesting that strength is a key functional feature that enables males to perform faster movements (but for shorter periods), associated with the demands of territoriality.     

Neuromuscular partitioning, architectural design, and myosin fiber types of the M. vastus lateralis of the llama (Lama glama)

The llama (Lama glama) is one of the few mammals of relatively large body size in which three fast myosin heavy chain isoforms (i.e., IIA, IIX, IIB) are extensively expressed in their locomotory muscles. This study was designed to gain insight into the morphological and functional organization of skeletal musculature in this peculiar animal model. The neuromuscular partitioning, architectural design, and myosin fiber types were systematically studied in the M. vastus lateralis of adult llamas (n = 15). Four nonoverlapping neuromuscular partitions or compartments were identified macroscopically (using a modified Sihler's technique for muscle depigmentation), although they did not conform strictly to the definitions of "neuromuscular compartments." Each neuromuscular partition was innervated by primary branches of the femoral nerve and was arranged within the muscle as paired partitions, two in parallel (deep-superficial compartmentalization) and the other two in-series (proximo-distal compartmentalization). These neuromuscular partitions of the muscle varied in their respective architectural designs (studied after partial digestion with diluted nitric acid) and myosin fiber type characteristics (identified immunohistochemically with specific anti-myosin monoclonal antibodies, then examined by quantitative histochemistry and image analysis). The deep partitions of the muscle had longer fibers, with lower angles of pinnation, and higher percentages of fast-glycolytic fibers than the superficial partitions of the muscle. These differences clearly suggest a division of labor in the whole M. vastus lateralis of llamas, with deep partitions exhibiting features well adapted for dynamic activities in the extension of stifle, whereas superficial portions seem to be related to the antigravitational role of the muscle in preserving the extension of the stifle during standing and stance phase of the stride. This peculiar structural and functional organization of the llama M. vastus lateralis does not confirm the generalized idea that deep muscles or the deepest portions within the same muscles somehow develop postural and/or low-intensity isometric functions. Rather, it suggests a primacy of architecture over intramuscular location in determining fiber type composition and hence division of labor within the muscle. A compartmentalization in the distribution of the three fast-subtype fibers (IIA, IIX, and IIB) also occurred, and this could also be relevant functionally, since these fiber types differed significantly in size (IIA < IIX < IIB), oxidative capacity (IIA > IIX > IIB), and capillarization (IIA = IIX > IIB). Furthermore, a typical spatial pattern in fiber type distribution was encountered in llama muscle (i.e., fiber types were consistently ranked in the order I --> IIA --> IIX --> IIB from the center to the periphery of fascicles), suggesting again peculiar and not well understood functional adaptations in these species.