A three-dimensional model of the rat hindlimb: musculoskeletal geometry and muscle moment arms

As a first step towards developing a dynamic model of the rat hindlimb, we measured muscle attachment and joint center coordinates relative to bony landmarks using stereophotogrammetry. Using these measurements, we analyzed muscle moment arms as functions of joint angle for most hindlimb muscles, and tested the hypothesis that postural change alone is sufficient to alter the function of selected muscles of the leg. We described muscle attachment sites as second-order curves. The length of the fit parabola and residual errors in the orthogonal directions give an estimate of muscle attachment sizes, which are consistent with observations made during dissection. We modeled each joint as a moving point dependent on joint angle; relative endpoint errors less than 7% indicate this method as accurate. Most muscles have moment arms with a large range across the physiological domain of joint angles, but their moment arms peak and vary little within the locomotion domain. The small variation in moment arms during locomotion potentially simplifies the neural control requirements during this phase. The moment arms of a number of muscles cross zero as angle varies within the quadrupedal locomotion domain, indicating they are intrinsically stabilizing. However, in the bipedal locomotion domain, the moment arms of these muscles do not cross zero and thus are no longer intrinsically stabilizing. We found that muscle function is largely determined by the change in moment arm with joint angle, particularly the transition from quadrupedal to bipedal posture, which may alter an intrinsically stabilizing arrangement or change the control burden.     

Cupiennius salei: biomechanical properties of the tibia–metatarsus joint and its flexing muscles

Hunting spiders are well adapted to fast locomotion. Space saving hydraulic leg extension enables leg segments, which consist almost soley of flexor muscles. As a result, the muscle cross sectional area is high despite slender legs. Considering these morphological features in context with the spider’s segmented C-shaped legs, these specifics might influence the spider’s muscle properties. Moreover, these properties have to be known for modeling of spider locomotion. Cupiennius salei (n = 5) were fixed in a metal frame allowing exclusive flexion of the tibia–metatarsus joint of the second leg (counted from anterior). Its flexing muscles were stimulated supramaximally using needle electrodes. Accounting for the joint geometry, the force–length and the force–velocity relationships were determined. The spider muscles produce 0.07 N cm maximum isometric moment (corresponding to 25 N/cm² maximum stress) at 160° tibia–metatarsus joint angle. When overextended to the dorsal limit at approximately 200°, the maximum isometric moments decrease to 72%, and, when flexed to the ventral hinge stop at 85°, they drop to 11%. The force–velocity relation shows the typical hyperbolic shape. The mean maximum shortening velocity is 5.7 optimum muscle lengths per second and the mean curvature (a/F _iso) of the Hill-function is 0.34. The spider muscle’s properties which were determined are similar to those of other species acting as motors during locomotion (working range, curvature of Hill hyperbola, peak power at the preferred speeds), but they are relatively slow. In conjunction with the low mechanical advantage (muscle lever/load arm), the arrangement of three considerably actuated joints in series may nonetheless enable high locomotion velocities.     

Locomotion in ornithischian dinosaurs: an assessment using three‐dimensional computational modelling

Ornithischian dinosaurs were primitively bipedal with forelimbs modified for grasping, but quadrupedalism evolved in the clade on at least three occasions independently. Outside of Ornithischia, quadrupedality from bipedal ancestors has only evolved on two other occasions, making this one of the rarest locomotory transitions in tetrapod evolutionary history. The osteological and myological changes associated with these transitions have only recently been documented, and the biomechanical consequences of these changes remain to be examined. Here, we review previous approaches to understanding locomotion in extinct animals, which can be broadly split into form–function approaches using analogy based on extant animals, limb‐bone scaling, and computational approaches. We then carry out the first systematic attempt to quantify changes in locomotor muscle function in bipedal and quadrupedal ornithischian dinosaurs. Using three‐dimensional computational modelling of the major pelvic locomotor muscle moment arms, we examine similarities and differences among individual taxa, between quadrupedal and bipedal taxa, and among taxa representing the three major ornithischian lineages (Thyreophora, Ornithopoda, Marginocephalia). Our results suggest that the ceratopsid Chasmosaurus and the ornithopod Hypsilophodon have relatively low moment arms for most muscles and most functions, perhaps suggesting poor locomotor performance in these taxa. Quadrupeds have higher abductor moment arms than bipeds, which we suggest is due to the overall wider bodies of the quadrupeds modelled. A peak in extensor moment arms at more extended hip angles and lower medial rotator moment arms in quadrupeds than in bipeds may be due to a more columnar hindlimb and loss of medial rotation as a form of lateral limb support in quadrupeds. We are not able to identify trends in moment arm evolution across Ornithischia as a whole, suggesting that the bipedal ancestry of ornithischians did not constrain the development of quadrupedal locomotion via a limited number of functional pathways. Functional anatomy appears to have had a greater effect on moment arms than phylogeny, and the differences identified between individual taxa and individual clades may relate to differences in locomotor performance required for living in different environments or for clade‐specific behaviours.     

Locomotor variation and bending regimes of capuchin limb bones

Primates are very versatile in their modes of progression, yet laboratory studies typically capture only a small segment of this variation. In vivo bone strain studies in particular have been commonly constrained to linear locomotion on flat substrates, conveying the potentially biased impression of stereotypic long bone loading patterns. We here present substrate reaction forces (SRF) and limb postures for capuchin monkeys moving on a flat substrate (“terrestrial”), on an elevated pole (“arboreal”), and performing turns. The angle between the SRF vector and longitudinal axes of the forearm or leg is taken as a proxy for the bending moment experienced by these limb segments. In both frontal and sagittal planes, SRF vectors and distal limb segments are not aligned, but form discrepant angles; that is, forces act on lever arms and exert bending moments. The positions of the SRF vectors suggest bending around oblique axes of these limb segments. Overall, the leg is exposed to greater moments than the forearm. Simulated arboreal locomotion and turns introduce variation in the discrepancy angles, thus confirming that expanding the range of locomotor behaviors studied will reveal variation in long bone loading patterns that is likely characteristic of natural locomotor repertoires. “Arboreal” locomotion, even on a linear noncompliant branch, is characterized by greater variability of force directions and discrepancy angles than “terrestrial” locomotion (significant for the forearm only), partially confirming the notion that life in trees is associated with greater variation in long bone loading. Directional changes broaden the range of external bending moments even further. Am J Phys Anthropol, 2009. © 2009 Wiley‐Liss, Inc.     

Modeling the effects of musculoskeletal geometry on scapulohumeral muscle moment arms and lines of action

Abstract

Biomechanical investigations examining shoulder function commonly observe a high degree of inter-individual variability in muscle activity and kinematic patterns during static and dynamic upper extremity exertions. Substantial differences in musculoskeletal geometry between individuals can alter muscle moment arms and lines of action that, theoretically, alter muscle activity and shoulder kinematics. The purposes of this research were to: (i) quantify model-predicted functional roles (moment arms, lines of action) of the scapulohumeral muscles, (ii) compare model predictions to experimental data in the literature, and (iii) evaluate sensitivity of muscle functional roles due to changes in muscle attachment locations using probabilistic modeling. Monte Carlo simulations were performed to iteratively adjust muscle attachment locations at the clavicle, scapula, and humerus of the Delft Shoulder and Elbow Model in OpenSim. Muscle moment arms and lines of action were quantified throughout arm elevation in the scapular plane. In general, model-predicted moment arms agreed well with the reviewed literature; however, notable inconsistencies were observed when comparing lines of action. Variability in moment arms and lines of action were muscle-specific, with 2 standard deviations in moment arm and line of actions as high as 25.8?mm and 18.8° for some muscles, respectively. Moment arms were particularly sensitive to changes in attachment site closest to the joint centre. Variations in muscle functional roles due to differences in musculoskeletal geometry are expected to require different muscle activity and movement patterns for upper extremity exertions.     

Locomotion pattern and trunk musculoskeletal architecture among Urodela

We comparatively examined the trunk musculature and prezygapophyseal angle of mid‐trunk vertebra in eight urodele species with different locomotive modes (aquatic Siren intermedia, Amphiuma tridactylum, Necturus maculosus and Andrias japonicus; semi‐aquatic Cynops pyrrhogaster, Cynops ensicauda; and terrestrial Hynobius nigrescens, Hynobius lichenatus and Ambystoma tigrinum). We found that the more terrestrial species were characterized by larger dorsal and abdominal muscle weight ratios compared with those of the more aquatic species, whereas muscle ratios of the lateral hypaxial musculature were larger in the more aquatic species. The lateral hypaxial muscles were thicker in the more aquatic species, whereas the M. rectus abdominis was more differentiated in the more terrestrial species. Our results suggest that larger lateral hypaxial muscles function for lateral bending during underwater locomotion in aquatic species. Larger dorsalis and abdominal muscles facilitate resistance against sagittal extension of the trunk, stabilization and support of the ventral contour line against gravity in terrestrial species. The more aquatic species possessed a more horizontal prezygapophyseal angle for more flexible lateral locomotion. In contrast, the more terrestrial species have an increasingly vertical prezygapophyseal angle to provide stronger column support against gravity. Thus, we conclude trunk structure in urodeles differs clearly according to their locomotive modes.     

Comparison of hind limb muscle mass in neonate and adult prosimian primates

Little ontogenetic data exist to indicate whether muscular organization of neonates reflects adult locomotion (e.g., leaping) or infant activities like clinging or the initial quadrupedal phase of locomotion that typifies most infant primates. In the present study, five species of primates with contrasting modes of locomotion were examined. Twenty-eight preserved neonatal and adult cadavers were studied by careful dissection of the hip, thigh, and leg muscles. Wet weights were taken of limb muscles after removal, and the muscles were combined into major functional groups (e.g., flexors, extensors) of each limb segment. Results demonstrate that the distribution of muscle mass within the thigh and within the leg are similar between neonates and adults for all species, with major groups varying by 5% or less in all but two age comparisons. Crural indices of the neonates are nearly identical to those of the adults, but leg/thigh muscle mass ratios were higher in the neonates. Species vary greatly in the percentage of adult limb segment muscle mass present in neonates, with Tarsius syrichta having the greatest percentage for all segments and two lemurids showing the least. These results primarily track differences in relative body mass at birth rather than developmental differences. The adaptive distribution of muscle, as discussed previously for adult prosimians, appears to be established at birth. Neonates of leaping species already have much larger quadriceps muscles than quadrupeds. Differences between large- and small-bodied leapers (e.g., pronounced superficial plantarflexor masses in tarsiers and pronounced deep plantarflexor masses in sifakas) also are present in neonates. Ratios of muscle mass over body mass are smaller in all neonates than in their adult counterparts, suggesting that the neonates are relatively poorly muscled, and that muscle mass must increase with positive allometry during growth.     

Dimensions and moment arms of the hind- and forelimb muscles of common chimpanzees (Pan troglodytes).

This paper supplies quantitative data on the hind- and forelimb musculature of common chimpanzees (Pan troglodytes) and calculates maximum joint moments of force as a contribution to a better understanding of the differences between chimpanzee and human locomotion. We dissected three chimpanzees, and recorded muscle mass, fascicle length, and physiological cross-sectional area (PCSA). We also obtained flexion/extension moment arms of the major muscles about the limb joints. We find that in the hindlimb, chimpanzees possess longer fascicles in most muscles but smaller PCSAs than are predicted for humans of equal body mass, suggesting that the adaptive emphasis in chimpanzees is on joint mobility at the expense of tension production. In common chimpanzee bipedalism, both hips and knees are significantly more flexed than in humans, necessitating muscles capable of exerting larger moments at the joints for the same ground force. However, we find that when subject to the same stresses, chimpanzee hindlimb muscles provide far smaller moments at the joints than humans, particularly the quadriceps and plantar flexors. In contrast, all forelimb muscle masses, fascicle lengths, and PCSAs are smaller in humans than in chimpanzees, reflecting the use of the forelimbs in chimpanzee, but not human, locomotion. When subject to the same stresses, chimpanzee forelimb muscles provide larger moments at the joints than humans, presumably because of the demands on the forelimbs during locomotion. These differences in muscle architecture and function help to explain why chimpanzees are restricted in their ability to walk, and particularly to run bipedally.     

Fiber‐type distribution of the perivertebral musculature in Ambystoma

Many salamanders locomote in aquatic and terrestrial environments. During swimming, body propulsion is solely produced by the axial musculature generating lateral undulations of the trunk and tail. During terrestrial locomotion, the trunk is oscillated laterally in a standing wave, and body propulsion is achieved by concerted trunk and limb muscle action. The goal of this study was to increase our knowledge of the functional morphology of the tetrapod trunk. We investigated the muscle‐fiber‐type distribution and the anatomical cross‐sectional area of all perivertebral muscles in Ambystoma tigrinum and A. maculatum. Muscle‐fiber‐type composition was determined in serial cross‐sections based on m‐ATPase activity. Five different body segments were investigated to test for cranio‐caudal changes along the trunk. The overall fiber‐type distribution was very similar between the species, but A. tigrinum had relatively larger muscles than A. maculatum, which may be related to its digging behavior. None of the perivertebral muscles possessed a homogeneous fiber‐type composition. The M. interspinalis showed a distinct layered organization and may function to ensure the integrity of the spine (local stabilization). The M. dorsalis trunci exhibited the plesiomorphic pattern for notochordates in having a distinct superficial layer of red and intermediate fibers, which covered the central white fibers; therefore, it is suggested to function as a mobilizer and a stabilizer of the trunk, but, may also be involved in modulating body stiffness. Similarly, the M. subvertebralis showed clear regionalizations, implying functional subunits that can stabilize and mobilize the trunk as well as modulate of body stiffness. Cranio‐caudally, neither the fiber‐type composition nor the a‐csa changed dramatically, possibly reflecting the need to perform well in both aquatic and terrestrial habitats. J. Morphol., 2010. © 2009 Wiley‐Liss, Inc.     

Muscle moment arm and normalized moment contributions as reference data for musculoskeletal elbow and wrist joint models

A geometric musculoskeletal model of the elbow and wrist joints was developed to calculate muscle moment arms throughout elbow flexion/extension, forearm pronation/supination, wrist flexion/extension and radial/ulnar deviation. Model moment arms were verified with data from cadaver specimen studies and geometric models available in the literature. Coefficients of polynomial equations were calculated for all moment arms as functions of joint angle, with special consideration to coupled muscles as a function of two joint angles. Additionally, a “normalized potential moment (NPM)” contribution index for each muscle across the elbow and wrist joints in four degrees-of-freedom was determined using each muscle's normalized physiological cross-sectional area (PCSA) and peak moment arm (MA). We hypothesize that (a) a geometric model of the elbow and wrist joints can represent the major attributes of MA versus joint angle from many literature sources of cadaver and model data and (b) an index can represent each muscle's normalized moment contribution to each degree-of-freedom at the elbow and wrist. We believe these data serve as a simple, yet comprehensive, reference for how the primary 16 muscles across the elbow and wrist contribute to joint moment and overall joint performance.     

Morphology and morphometrics of the appendicular musculature in geckoes with different locomotor habits (Lepidosauria)

 In this study a ground-dwelling (Eublepharis macularius) and a highly specialised climbing (Gekko gecko) lizard were chosen as study objects. The fore- and hindlimbs of two individuals of each species were dissected, and muscle masses, mean fibre lengths, cross-sectional areas and moment arms were determined. Special attention was paid to general muscle architecture (origin, insertion, fibre orientation, etc.) and pennation angles. Using these variables (cross sectional areas and moment arms), maximal moments exertable across the shoulder/hip, elbow/knee and wrist/ankle were calculated for both species. In accordance with the biomechanical predictions related to the preferred locomotor substrate of each species (i.e. level running for E. macularius and climbing for G. gecko), the results of this study indicate that climbers such as G. gecko generally possess powerful retractor muscles crossing the shoulder and hip joints. Additionally, the specialised climber is able to exert higher flexion moments across the elbow, which prevents the animals from falling backwards. However, G. gecko appears to be constrained in its ankle extension capabilities by the presence of the adhesive toe pads. The level-running species, on the other hand, shows a relatively stronger development of the extensor muscles in the lower limbs, allowing these lizards to run in an erect posture. In general, both species show large similarities on a gross morphological level as expected when considering their phylogenetic relatedness. Adaptations to their preferred locomotor substrate only become apparent when considering the functional properties (i.e. joint moments) of the appendicular musculature. Accepted: 30 November 1998     

Sensitivity of model predictions of muscle function to changes in moment arms and muscle-tendon properties: a Monte-Carlo analysis

Hill-type muscle models are commonly used in musculoskeletal models to estimate muscle forces during human movement. However, the sensitivity of model predictions of muscle function to changes in muscle moment arms and muscle-tendon properties is not well understood. In the present study, a three-dimensional muscle-actuated model of the body was used to evaluate the sensitivity of the function of the major lower limb muscles in accelerating the whole-body center of mass during gait. Monte-Carlo analyses were used to quantify the effects of entire distributions of perturbations in the moment arms and architectural properties of muscles. In most cases, varying the moment arm and architectural properties of a muscle affected the torque generated by that muscle about the joint(s) it spanned as well as the torques generated by adjacent muscles. Muscle function was most sensitive to changes in tendon slack length and least sensitive to changes in muscle moment arm. However, the sensitivity of muscle function to changes in moment arms and architectural properties was highly muscle-specific; muscle function was most sensitive in the cases of gastrocnemius and rectus femoris and insensitive in the cases of hamstrings and the medial sub-region of gluteus maximus. The sensitivity of a muscle's function was influenced by the magnitude of the muscle's force as well as the operating region of the muscle on its force-length curve. These findings have implications for the development of subject-specific models of the human musculoskeletal system.     

Biarticular hip extensor and knee flexor muscle moment arms of the feline hindlimb

Moment arms are important for understanding muscular behavior and for calculating internal muscle forces in musculoskeletal simulations. Biarticular muscles cross two joints and have moment arms that depend on the angle of both joints the muscles cross. The tendon excursion method was used to measure the joint angle-dependence of hamstring (biceps femoris, semimembranosus and semitendinosus) moment arm magnitudes of the feline hindlimb at the knee and hip joints. Knee angle influenced hamstring moment arm magnitudes at the hip joint; compared to a flexed knee joint, the moment arm for semimembranosus posterior at the hip was at most 7.4 mm (25%) larger when the knee was extended. On average, hamstring moment arms at the hip increased by 4.9 mm when the knee was more extended. In contrast, moment arm magnitudes at the knee varied by less than 2.8 mm (mean=1.6 mm) for all hamstring muscles at the two hip joint angles tested. Thus, hamstring moment arms at the hip were dependent on knee position, while hamstring moment arms at the knee were not as strongly associated with relative hip position. Additionally, the feline hamstring muscle group had a larger mechanical advantage at the hip than at the knee joint.     

How muscles accommodate movement in different physical environments: aquatic vs. terrestrial locomotion in vertebrates.

Representatives of nearly all vertebrate classes are capable of coordinated movement through aquatic and terrestrial environments. Though there are good data from a variety of species on basic patterns of muscle recruitment during locomotion in a single environment, we know much less about how vertebrates use the same musculoskeletal structures to accommodate locomotion in physically distinct environments. To address this issue, we have gathered data from a broad range of vertebrates that move successfully through water and across land, including eels, toads, turtles and rats. Using high-speed video in combination with electromyography and sonomicrometry, we have quantified and compared the activity and strain of individual muscles and the movements they generate during aquatic vs. terrestrial locomotion. In each focal species, transitions in environment consistently elicit alterations in motor output by major locomotor muscles, including changes in the intensity and duration of muscle activity and shifts in the timing of activity with respect to muscle length change. In many cases, these alterations likely change the functional roles played by muscles between aquatic and terrestrial locomotion. Thus, a variety of forms of motor plasticity appear to underlie the ability of many species to move successfully through different physical environments and produce diverse behaviors in nature.     

Development of an anatomically based whole‐body musculoskeletal model of the Japanese macaque (Macaca fuscata)

We constructed a three‐dimensional whole‐body musculoskeletal model of the Japanese macaque (Macaca fuscata) based on computed tomography and dissection of a cadaver. The skeleton was modeled as a chain of 20 bone segments connected by joints. Joint centers and rotational axes were estimated by joint morphology based on joint surface approximation using a quadric function. The path of each muscle was defined by a line segment connecting origin to insertion through an intermediary point if necessary. Mass and fascicle length of each were systematically recorded to calculate physiological cross‐sectional area to estimate the capacity of each muscle to generate force. Using this anatomically accurate model, muscle moment arms and force vectors generated by individual limb muscles at the foot and hand were calculated to computationally predict muscle functions. Furthermore, three‐dimensional whole‐body musculoskeletal kinematics of the Japanese macaque was reconstructed from ordinary video sequences based on this model and a model‐based matching technique. The results showed that the proposed model can successfully reconstruct and visualize anatomically reasonable, natural musculoskeletal motion of the Japanese macaque during quadrupedal/bipedal locomotion, demonstrating the validity and efficacy of the constructed musculoskeletal model. The present biologically relevant model may serve as a useful tool for comprehensive understanding of the design principles of the musculoskeletal system and the control mechanisms for locomotion in the Japanese macaque and other primates. Am J Phys Anthropol, 2009. © 2008 Wiley‐Liss, Inc.     

Rotational moment arms of the medial hamstrings and adductors vary with femoral geometry and limb position: implications for the treatment of internally rotated gait

Persons with cerebral palsy frequently walk with a crouched, internally rotated gait. Spastic medial hamstrings or adductors are presumed to contribute to excessive hip internal rotation in some patients; however, the capacity of these muscles to produce internal rotation has not been adequately investigated. The purpose of this study was to determine the hip rotation moment arms of the medial hamstrings and adductors in persons with femoral anteversion deformities who walk with a crouched, internally rotated gait. A musculoskeletal model with a "deformable" femur was developed. This model was used, in conjunction with kinematic data obtained from gait analysis, to calculate the muscle moment arms for combinations of joint angles and anteversion deformities exhibited by 21 subjects with cerebral palsy and excessive hip internal rotation. We found that the semimembranosus, semitendinosus, and gracilis muscles in our model had negligible or external rotation moment arms when the hip was internally rotated or the knee was flexed -- the body positions assumed by the subjects during walking. When the femur was excessively anteverted, the rotational moment arms of the adductor brevis, adductor longus, pectineus, and proximal compartments of the adductor magnus in our model shifted toward external rotation. These results suggest that neither the medial hamstrings nor the adductors are likely to contribute substantially to excessive internal rotation of the hip and that other causes of internal rotation should be considered when planning treatments for these patients.     

Influence of vibration on the excitability of spinal α-motoneurons under static conditions and during evoked stepping in humans

The excitability of spinal α-motoneurons in healthy humans was investigated with vibrostimulation (20–60 Hz) applied to different groups of muscles both under stationary conditions and during vibration-evoked stepping movements with leg suspension. In 15 subjects, the H-reflex amplitude was compared under the conditions of vibration of the left leg quadriceps femoris (QFM) or biceps femoris (BFM) muscle, as well as under the conditions of vibration of the contralateral, motionless leg QFM muscle in three spatial positions of the body: upright, supine, and lying on the side with the left leg suspended. Under dynamic conditions, the H-reflex value was compared during evoked and voluntary steppings at eight intervals of the step cycle. In all body positions, the vibration of each ipsilateral leg muscle caused a significant H-reflex suppression, this suppression being more prominent under the air-stepping conditions. The vibration of the contralateral leg QFM had weak influence on the H-reflex amplitude. In seven subjects, the vibration of the ipsilateral and contralateral leg muscles generated stepping movements. During vibration-evoked air-stepping, the H-reflex had different amplitudes in different phases of the step cycle. At the same time, the differences between responses under voluntary and involuntary stepping conditions were revealed only in the step cycle phase corresponding to the stance phase. Thus, the different degrees of the H-reflex suppression by vibration in different spatial positions of the body seem to depend on the summary afferent inflows to the spinal cord interneurons involved in the regulation of locomotion and posture. Apparently, an increase in the spinal cord neuronal excitability, which is necessary for activating locomotor automatism under the leg unloading conditions, occurs during evoked air-stepping in the swing phase.     

Design of the predatory legs of water bugs (Hemiptera: Nepidae,Naucoridae, Notonectidae,Gerridae)

Functional comparative morphology of predatory legs in five species of water bugs (Ilyocoris cimicoides, Nepa cinerea, Ranatra linearis, Notonecta glauca, and Gerris lacustris) has been investigatd adn the following peculiarities of leg design were revealed.

• 1 Subcoxal articulation may be monoaxial (G. lacustris, N. glauca), or, in contrast to walking leg type, biaxial (N. cinerea, R. linearis, I. cimicoides); the first axis is oriented along the coxa (torsion axis), the second one is perpendicular to the first (non-torsion axis).
• 2 In contrast to walking leg type, which is characterized by cross suspension of the axis of coxal rotation in thoracal skeleton, this axis in G. lacustris is placed vertically. Non-torsion coxal axis in R. linearis is oriented strongly transversal. This axis directs the leg strike forward.
• 3 Legs in the majority of species are planar: Torsion axes of the coxa, femur, and tibia are placed in the same plane. Axes of rotation of consequent joints in I. cimicoides are reciprocally sloped. Therefore, the end of the leg outlines the spiral trajectory, when all angles of joints are opening (closing). This is an adaptation for clinging to the stems of water plants.
• 4 Passive adduction of the femur in the trochanter-femoral joint in N. glauca allows it to go around protuberances of the body wall, when the leg is sliding along them; recurrent femur movement during releasing from the obstacele is active due to the rt.fe muscle.
• 5 Only R. linearis has predatory legs, which permit the high-speed pursuit of potential prey; other species realize this function using the swimming legs, whereas the forelegs are used for the manipulation movements.
• 6 Muscle arrangement in the prothorax of different species reflects both leg construction and constructional constraints of body design. Powerful flexor muscles (co1, co2, co3, co5, fl.ti, et.ti in R. linearis; fl.ta, fl.ti in N. glauca; fl.ti in I. cimicoides) have long tendons and short muscle bundles, which originate on the leg wall. As a result, the powerful force is developed along the muscle tendon.
• 7 Some features of the predatory leg are common for the species studies: elongation of coxae, thickening of femora, and increase of the degree of junction of tibia and tarsus. The muscles, which move the distal segment of the leg, are reinforced and the sclerite of the fl.ti tendon is enlarged. The joint angle of the distal segment is increased to 120°. © 1995 Wiley-Liss, Inc.

     

Scaling of peak moment arms of elbow muscles with upper extremity bone dimensions.

It is often assumed that moment arms scale with size and can be normalized by body segment lengths or limb circumferences. However, quantitative scaling relationships between moment arms and anthropometric dimensions are generally not available. We hypothesized that peak moment arms of the elbow flexor and extensor muscles scale with the shorter distance (D(s)) between the elbow flexion axis and a muscle's origin and insertion. To test this hypothesis, we estimated moment arms of six muscles that cross the elbow, digitized muscle attachment sites and bone surface geometry, and estimated the location of the elbow flexion axis in 10 upper extremity cadaveric specimens which ranged in size from a 5'0" female to a 6'4" male. D(s) accurately reflected the differences in peak moment arms across different muscles, explaining 93-99% of the variation in peaks between muscles in the same specimen. D(s) also explained between 55% and 88% of the interspecimen variation in peak moment arms for brachioradialis, biceps, and ECRL. Triceps peak moment arm was significantly correlated to the anterior-posterior dimension of the ulna measured at the olecranon (r(2)=0.61, p=0.008). Radius length provides a good measure of the interspecimen variation in peaks for brachioradialis, biceps, and ECRL. However, bone lengths were not significantly correlated to triceps moment arm or anterior-posterior bone dimensions. This work advances our understanding of the variability and scaling dimensions for elbow muscle moment arms across subjects of different sizes.     

Moment arms of the human neck muscles in flexion, bending and rotation

There is a paucity of data available for the moment arms of the muscles of the human neck. The objective of the present study was to measure the moment arms of the major cervical spine muscles in vitro. Experiments were performed on five fresh-frozen human head-neck specimens using a custom-designed robotic spine testing apparatus. The testing apparatus replicated flexion-extension, lateral bending and axial rotation of each individual intervertebral joint in the cervical spine while all other joints were kept immobile. The tendon excursion method was used to measure the moment arms of 30 muscle sub-regions involving 13 major muscles of the neck about all three axes of rotation of each joint for the neutral position of the cervical spine. Significant differences in the moment arm were observed across sub-regions of individual muscles and across the intervertebral joints spanned by each muscle (p<0.05). Overall, muscle moment arms were larger in flexion-extension and lateral bending than in axial rotation, and most muscles had prominent moment arms in at least 2 out of the 3 joint motions investigated. This study emphasizes the importance of detailed representation of a muscle's architecture in prediction of its torque capacity about the individual joints of the cervical spine. The dataset produced may be useful in developing and validating computational models of the human neck.