Back muscle function during bipedal walking in chimpanzee and gibbon: implications for the evolution of human locomotion

The evolution of erect posture and locomotion continues to be a major focus of interest among paleoanthropologists and functional morphologists. To date, virtually all of our knowledge about the functional role of the back muscles in the evolution of bipedalism is based on human experimental data. In order to broaden our evolutionary perspective on the vertebral region, we have undertaken an electromyographic (EMG) analysis of three deep back muscles (multifidus, longissimus thoracis, iliocostalis lumborum) in the chimpanzee (Pan troglodytes) and gibbon (Hylobates lar) during bipedal walking. The recruitment patterns of these three muscles seen in the chimpanzee closely parallel those observed in the gibbon. The activity patterns of multifidus and longissimus are more similar to each other than either is to iliocostalis. Iliocostalis recruitment is clearly related to contact by the contralateral limb during bipedal walking in both species. It is suggested that in both the chimpanzee and gibbon, multifidus controls trunk movement primarily in the sagittal plane, iliocostalis responds to and adjusts movement in the frontal plane, while longissimus contributes to both of these functions. In many respects, the activity patterns shared by the chimpanzee and gibbon are quite consistent with recent human experimental data. This suggests a basic similarity in the mechanical constraints placed on the back during bipedalism among these three hominoids. Thus, the acquisition of habitual bipedalism in humans probably involved not so much a major change in back muscle action or function, but rather an improvement in the mechanical advantages and architecture of these muscles.     

Isoform diversity of giant proteins in relation to passive and active contractile properties of rabbit skeletal muscles

The active and passive contractile performance of skeletal muscle fibers largely depends on the myosin heavy chain (MHC) isoform and the stiffness of the titin spring, respectively. Open questions concern the relationship between titin-based stiffness and active contractile parameters, and titin's importance for total passive muscle stiffness. Here, a large set of adult rabbit muscles (n = 37) was studied for titin size diversity, passive mechanical properties, and possible correlations with the fiber/MHC composition. Titin isoform analyses showed sizes between approximately 3300 and 3700 kD; 31 muscles contained a single isoform, six muscles coexpressed two isoforms, including the psoas, where individual fibers expressed similar isoform ratios of 30:70 (3.4:3.3 MD). Gel electrophoresis and Western blotting of two other giant muscle proteins, nebulin and obscurin, demonstrated muscle type-dependent size differences of < or =70 kD. Single fiber and single myofibril mechanics performed on a subset of muscles showed inverse relationships between titin size and titin-borne tension. Force measurements on muscle strips suggested that titin-based stiffness is not correlated with total passive stiffness, which is largely determined also by extramyofibrillar structures, particularly collagen. Some muscles have low titin-based stiffness but high total passive stiffness, whereas the opposite is true for other muscles. Plots of titin size versus percentage of fiber type or MHC isoform (I-IIB-IIA-IID) determined by myofibrillar ATPase staining and gel electrophoresis revealed modest correlations with the type I fiber and MHC-I proportions. No relationships were found with the proportions of the different type II fiber/MHC-II subtypes. Titin-based stiffness decreased with the slow fiber/MHC percentage, whereas neither extramyofibrillar nor total passive stiffness depended on the fiber/MHC composition. In conclusion, a low correlation exists between the active and passive mechanical properties of skeletal muscle fibers. Slow muscles usually express long titin(s), predominantly fast muscles can express either short or long titin(s), giving rise to low titin-based stiffness in slow muscles and highly variable stiffness in fast muscles. Titin contributes substantially to total passive stiffness, but this contribution varies greatly among muscles.     

Passive tension of rat skeletal soleus muscle fibers: effects of unloading conditions.

In this work we studied changes in passive elastic properties of rat soleus muscle fibers subjected to 14 days of hindlimb unloading (HU). For this purpose, we investigated the titin isoform expression in soleus muscles, passive tension-fiber strain relationships of single fibers, and the effects of the thick filament depolymerization on passive tension development. The myosin heavy chain composition was also measured for all fibers studied. Despite a slow-to-fast transformation of the soleus muscles on the basis of their myosin heavy chain content, no modification in the titin isoform expression was detected after 14 days of HU. However, the passive tension-fiber strain relationships revealed that passive tension of both slow and fast HU soleus fibers increased less steeply with sarcomere length than that of control fibers. Gel analysis suggested that this result could be explained by a decrease in the amount of titin in soleus muscle after HU. Furthermore, the thick filament depolymerization was found to similarly decrease passive tension in control and HU soleus fibers. Taken together, these results suggested that HU did not change titin isoform expression in the soleus muscle, but rather modified muscle stiffness by decreasing the amount of titin.     

Characterization of the passive mechanical properties of spine muscles across species

Passive mechanical properties differ between muscle groups within a species. Altered functional demands can also shift the passive force-length relationship. The extent that passive mechanical properties differ within a muscle group (e.g. spine extensors) or between homologous muscles of different species is unknown. It was hypothesized that multifidus, believed to specialize in spine stabilization, would generate greater passive tensile stresses under isometric conditions than erector spinae, which have more generalized functions of moving and stabilizing the spine; observing greater multifidus moduli in different species would strengthen this hypothesis. Permeabilized fibre bundles (n = 337) from the multifidus and erector spinae of mice, rats, and rabbits were mechanically tested. A novel logistic function was fit to the experimental data to fully characterize passive stress and modulus. Species had the greatest effect on passive muscle parameters with mice having the largest moduli at all lengths. Rats generated less passive stress than rabbits due to a shift of the passive force-length relationship towards longer muscle lengths. Rat multifidus generated slightly greater stresses than erector spinae, but no differences were observed between mouse muscles. The secondary objective was to determine the parameters required to simulate the passive force-length relationship. Experimental data were compared to the passive muscle model in OpenSim. The default OpenSim model, optimized for hindlimb muscles, did not fit any of the spine muscles tested; however, the model could accurately simulate experimental data after adjusting the input parameters. The optimal parameters for modelling the passive force-length relationships of spine muscles in OpenSim are presented.     

Epaxial Musculature in Armadillos,Sloths, and Opossums: Functional Significance and Implications for the Evolution of Back Muscles in the Xenarthra

To investigate the evolution of xenarthran epaxial muscles, fresh specimens of the North American Common long-nosed armadillo Dasypus novemcinctus and of a marsupial, the Virginia opossum Didelphis virginiana, were dissected. Data from one fixed specimen of a two-toed sloth Choloepus didactylus were also used for comparison, because it is a xenarthran exhibiting a highly derived locomotor mode. The opossum was used to represent a more generalized mammalian condition. Each of the three mammalian epaxial muscle groups, the iliocostalis, longissimus dorsi, and transversospinalis, was removed and its mass was determined. All data were corrected for body mass and length. Unpaired, one-tailed t-tests showed the average mass of the iliocostalis and transversospinalis of Dasypus to be significantly larger than the mass of the same muscles in Didelphis, whereas the average mass of the longissimus dorsi was not statistically different between the two species. In agreement with pronounced lateral bending and de-emphasized dorso-ventral flexion and extension, Choloepus also had a relatively large iliocostalis and small longissimus. Our limited data suggest that this condition was inherited from non-arboreal and probably digging early xenarthrans. We believe the relatively larger iliocostalis and transversospinalis muscles in Dasypus can be attributed to the need to provide vertical stabilization of the trunk and resist lateral reaction forces generated by digging. Thus, for Xenarthra it represents a synapomorphy linked to adaptations for fossoriality.     

Passive tension and stiffness of vertebrate skeletal and insect flight muscles: the contribution of weak cross-bridges and elastic filaments.

Tension and dynamic stiffness of passive rabbit psoas, rabbit semitendinosus, and waterbug indirect flight muscles were investigated to study the contribution of weak-binding cross-bridges and elastic filaments (titin and minititin) to the passive mechanical behavior of these muscles. Experimentally, a functional dissection of the relative contribution of actomyosin cross-bridges and titin and minititin was achieved by 1) comparing mechanically skinned muscle fibers before and after selective removal of actin filaments with a noncalcium-requiring gelsolin fragment (FX-45), and 2) studying passive tension and stiffness as a function of sarcomere length, ionic strength, temperature, and the inhibitory effect of a carboxyl-terminal fragment of smooth muscle caldesmon. Our data show that weak bridges exist in both rabbit skeletal muscle and insect flight muscle at physiological ionic strength and room temperature. In rabbit psoas fibers, weak bridge stiffness appears to vary with both thin-thick filament overlap and with the magnitude of passive tension. Plots of passive tension versus passive stiffness are multiphasic and strikingly similar for these three muscles of distinct sarcomere proportions and elastic proteins. The tension-stiffness plot appears to be a powerful tool in discerning changes in the mechanical behavior of the elastic filaments. The stress-strain and stiffness-strain curves of all three muscles can be merged into one, by normalizing strain rate and strain amplitude of the extensible segment of titin and minititin, further supporting the segmental extension model of resting tension development.     

Lumbar muscles recruitment during resistance exercise for upper limbs

The aim of this study was to evaluate the EMG activity of lumbar multifidus (MU), longissimus thoracis (LT) and iliocostalis (IC) muscles during an upper limb resistance exercise (biceps curl). Ten healthy males performed maximal voluntary isometric contraction (MVC) of the trunk extensors, after this, the biceps curl exercise was executed at 25%, 30%, 35% and 40% one repetition maximum during 1 min, with 10 min rest between them. EMG root mean square (RMS) and median frequency (MFreq) were calculated for each lifting and lowering of the bar during the exercise bouts, to calculate slopes and intercepts. The results showed increases in the RMS and decreases in the MFreq slopes. RMS slopes were no different between muscles, indicating similar fatigue process along the exercise irrespective of the load level. MU and LT presented higher RMS irrespective of the load level, which can be related to the specific function during the standing position. On the other hand, IC and MU presented higher MFreq intercepts compared to LT, demonstrating possible differences in the muscle fiber conduction velocity of these muscles. These findings suggest that trunk muscles are differently activate during upper limb exercises, and the fatigue process affects the lumbar muscles similarly.     

Anticipatory postural adjustments to arm movement reveal complex control of paraspinal muscles in the thorax

Anatomical and empirical data suggest that deep and superficial muscles may have different functions for thoracic spine control. This study investigated thoracic paraspinal muscle activity during anticipatory postural adjustments associated with arm movement. Electromyographic (EMG) recordings were made from the right deep (multifidus/rotatores) and superficial (longissimus) muscles at T5, T8, and T11 levels using fine-wire electrodes. Ten healthy participants performed fast unilateral and bilateral flexion and extension arm movements in response to a light. EMG amplitude was measured during 25 ms epochs for 150 ms before and 400 ms after deltoid EMG onset. During arm flexion movements, multifidus and longissimus had two bursts of activity, one burst prior to deltoid and a late burst. With arm extension both muscles were active in a single burst after deltoid onset. There was differential activity with respect to direction of trunk rotation induced by arm movement. Right longissimus was most active with left arm movements and right multifidus was most active with right arm movements. All levels of the thorax responded similarly. We suggest that although thoracic multifidus and longissimus function similarly to control sagittal plane perturbations, these muscles are differentially active with rotational forces on the trunk.     

Evaluation of measurement strategies to increase the reliability of EMG indices to assess back muscle fatigue and recovery.

The purpose of this study was to assess different measurement strategies to increase the reliability of different electromyographic (EMG) indices developed for the assessment of back muscle impairments. Forty male volunteers (20 controls and 20 chronic low back pain patients) were assessed on three sessions at least 2 days apart within 2 weeks. Surface EMG signals were recorded from four pairs (bilaterally) of back muscles (multifidus at the L5 level, iliocostalis lumborum at L3, and longissimus at L1 and T10) while the subjects performed, in a static dynamometer, two static trunk extension tasks at 75% of the maximal voluntary contraction separated by a 60 s rest period: (1) a 30 s fatigue task and (2) a 5 s recovery task. Different EMG indices (based on individual muscles or averaged across bilateral homologous muscles or across all muscles) were computed to evaluate muscular fatigue and recovery. Intra-class correlation coefficient (ICC) and standard error of measurement (SEM) in percentage of the grand mean were calculated for each EMG variable. Reliable EMG indices are achieved for both healthy and chronic low back pain subjects when (1) electrodes are positioned on medial back muscles (multifidus at the L5 level and longissimus at L1) and (2) measures are averaged across bilateral muscles and/or across two fatigue tests performed within a session. The most reliable EMG indices were the bilateral average of medial back muscles (ICC range: 0.68-0.91; SEM range: 5-35%) and the average of all back muscles (ICC range: 0.77-0.91; SEM range: 5-30%). The averaging of measures across two fatigue tests is predicted to increase the reliability by about 13%. With regards to EMG indices of fatigue, the identification of the most fatigable muscle also lead to satisfactory results (ICC range: 0.74-0.79; SEM range: 21-26%). The assessment of back muscle impairments through EMG analysis necessitates the use of multiple electrodes to achieve reliable results.     

Removal of immunoglobulin-like domains from titin’s spring segment alters titin splicing in mouse skeletal muscle and causes myopathy

Titin is a molecular spring that determines the passive stiffness of muscle cells. Changes in titin’s stiffness occur in various myopathies, but whether these are a cause or an effect of the disease is unknown. We studied a novel mouse model in which titin’s stiffness was slightly increased by deleting nine immunoglobulin (Ig)-like domains from titin’s constitutively expressed proximal tandem Ig segment (IG KO). KO mice displayed mild kyphosis, a phenotype commonly associated with skeletal muscle myopathy. Slow muscles were atrophic with alterations in myosin isoform expression; functional studies in soleus muscle revealed a reduced specific twitch force. Exon expression analysis showed that KO mice underwent additional changes in titin splicing to yield smaller than expected titin isoforms that were much stiffer than expected. Additionally, splicing occurred in the PEVK region of titin, a finding confirmed at the protein level. The titin-binding protein Ankrd1 was highly increased in the IG KO, but this did not play a role in generating small titin isoforms because titin expression was unaltered in IG KO mice crossed with Ankrd1-deficient mice. In contrast, the splicing factor RBM20 (RNA-binding motif 20) was also significantly increased in IG KO mice, and additional differential splicing was reversed in IG KO mice crossed with a mouse with reduced RBM20 activity. Thus, increasing titin’s stiffness triggers pathological changes in skeletal muscle, with an important role played by RBM20.     

Electromyography of back muscles during quadrupedal and bipedal walking in primates

Despite the extensive electromyographic research that has addressed limb muscle function during primate quadrupedalism, the role of the back muscles in this locomotor behavior has remained undocumented. We report here the results of an electromyographic (EMG) analysis of three intrinsic back muscles (multifidus, longissimus, and iliocostalis) in the baboon (Papio anubis), chimpanzee (Pan troglodytes), and orangutan (Pongo pygmaeus) during quadrupedal walking. The recruitment patterns of these three back muscles are compared to those reported for the same muscles during nonprimate quadrupedalism. In addition, the function of the back muscles during quadrupedalism and bipedalism in the two hominoids is compared. Results indicate that the back muscles restrict trunk movements during quadrupedalism by contracting with the touchdown of one or both feet, with more consistent activity associated with touchdown of the contralateral foot. Moreover, despite reported differences in their gait preferences and forelimb muscle EMG patterns, primates and nonprimate mammals recruit their back muscles in an essentially similar fashion during quadrupedal walking. These quadrupedal EMG patterns also resemble those reported for chimpanzees, gibbons and humans (but not orangutans) walking bipedally. The fundamental similarity in back muscle function across species and locomotor behaviors is consistent with other data pointing to conservatism in the evolution of the neural control of tetrapod limb movement, but does not preclude the suggestion (based on forelimb muscle EMG and spinal lesion studies) that some aspects of primate neural circuitry are unique. © 1994 Wiley-Liss, Inc.     

Spinal loads and trunk muscles forces during level walking – A combined in vivo and in silico study on six subjects

During level walking, lumbar spine is subjected to cyclic movements and intricate loading of the spinal discs and trunk musculature. This study aimed to estimate the spinal loads (T12–S1) and trunk muscles forces during a complete gait cycle.Six men, 24–33 years walk barefoot at self-selected speed (4–5 km/h). 3D kinematics and ground reaction forces were recorded using a motion capturing system and two force plates, implemented in an inverse dynamic musculoskeletal model to predict the spinal loads and trunk muscles forces. Additionally, the sensitivity of the intra-abdominal pressure and lumbar segment rotational stiffness was investigated.Peak spinal loads and trunk muscle forces were between the gait instances of heel strike and toe off. In L4–L5 segment, sensitivity analysis showed that average peak compressive, antero-posterior and medio-lateral shear forces were 130–179%, 2–15% and 1–6%, with max standard deviation (±STD) of 40%, 6% and 3% of the body weight. Average peak global muscles forces were 24–55% (longissimus thoracis), 11–23% (iliocostalis thoracis), 12–16% (external oblique), 17–25% (internal oblique) and 0–8% (rectus abdominus) of body weight whereas, the average peak local muscles forces were 11–19% (longissimus lumborum), 14–31% (iliocostalis lumborum) and 12–17% (multifidus). Maximum ± STD of the global and local muscles forces were 13% and 8% of the body weight.Large inter-individual differences were found in peak compressive and trunk muscles forces whereas the sensitivity analysis also showed a substantial variation.     

Intramuscular morphology and tendon geometry of the epaxial swimming muscles of dolphins

The dolphin upstroke is powered primarily by the m. multifidus and m. longissimus—robust muscles that insert serially along the vertebral column. Intramuscular and tendon morphology coupled with kinematic data are used to hypothesize regionally specific functions of these muscles. Both muscles develop equivalent forces (approximately 2kN) in the region of the thoraco-lumbar spine, but transmit those forces to different regions of the vertebral column. The m. multifidus transmits the majority of its force locally to the thoraco-lumar spine, a region of the body that undergoes no measurable bending. The action of the m. multifidus appears to be to stiffen its deep tendon of insertion, forming a temporary skeletal element for the m. longissimus. The m. longissimus transmits the majority of its force to the caudal spine, by way of a novel interaction between its insertional tendons and the subdermal connective tissue sheath. This insertional pattern confers torsional stiffness on the caudal peduncle, and allows the m. longissimus to do relatively more work (118 J) than if it had a typical mammalian insertional pattern. The caudal extension of the m. multifidus does 12 J of work on the caudal spine during an upstroke. The caudal extension of the m. longissimus transmits its force to the vertebrae in the caudal flukes and is the only epaxial muscle that acts to control the angle of attack of the fluke blade.     

Expression of titin isoforms in red and white muscle fibres of carp (Cyprinus carpio L.) exposed to different sarcomere strains during swimming

Titin (also known as connectin) is a striated-muscle-specific protein that spans the distance between the Z- and M-lines of the sarcomere. The elastic segment of the titin molecule in the I-band is thought to be responsible for developing passive tension and for maintaining the central position of thick filaments in contracting sarcomeres. Different muscle types express isoforms of titin that differ in their molecular mass. To help to elucidate the relation between the occurrence of titin isoforms and the functional properties of different fibre types, we investigated the presence of different titin isoforms in red and white fibres of the axial muscles of carp. Gel electrophoresis of single fibres revealed that the molecular mass of titin was larger in red than in white fibres. Fibres from anterior and posterior axial muscles were also compared. For both white and red fibres the molecular mass of titin in posterior muscle fibres was larger than in anterior muscle fibres. Thus, the same fibre type can express different titin isoforms depending on its location along the body axis. The contribution of titin to passive tension and stiffness of red anterior and posterior fibres was also determined. Single fibres were skinned and the sarcomere length dependencies of passive tension and passive stiffness were determined. Measurements were made before and after extracting thin and thick filaments using relaxing solutions with 0.6 mol · l⁻¹ KCl and 1 mol · l⁻¹ KI. Tension and stiffness measured before extraction were assumed to result from both titin and intermediate filaments, and tension after extraction from only intermediate filaments. Compared to mammalian skeletal muscle, intermediate filaments developed high levels of tension and stiffness in both posterior and anterior fibres. The passive tension-sarcomere length curve of titin increased more steeply in red anterior fibres than in red posterior fibres and the curve reached a plateau at a shorter sarcomere length. Thus, the smaller titin isoform of anterior fibres results in more passive tension and stiffness for a given sarcomere strain. During continuous swimming, red fibres are exposed to larger changes in sarcomere strain than white fibres, and posterior fibres to larger changes in strain than anterior fibres. We propose that sarcomere strain is one of the functional parameters that modulates the expression of different titin isoforms in axial muscle fibres of carp. Accepted: 7 May 1997     

Titin (Visco-) Elasticity in Skeletal Muscle Myofibrils

Titin is the third most abundant protein in sarcomeres and fulfills a number of mechanical and signaling functions. Specifically, titin is responsible for most of the passive forces in sarcomeres and the passive visco-elastic behaviour of myofibrils and muscles. It has been suggested, based on mechanical testing of isolated titin molecules, that titin is an essentially elastic spring if Ig domain un/refolding is prevented either by working at short titin lengths, prior to any unfolding of Ig domains, or at long sarcomere (and titin) lengths when Ig domain un/refolding is effectively prevented. However, these properties of titin, and by extension of muscles, have not been tested with titin in its natural structural environment within a sarcomere. The purpose of this study was to gain insight into the Ig domain un/refolding kinetics and test the idea that titin could behave essentially elastically at any sarcomere length by preventing Ig domain un/refolding during passive stretch-shortening cycles. Although not completely successful, we demonstrate here that titin’s visco-elastic properties appear to depend on the Ig domain un/refolding kinetics and that indeed, titin (and thus myofibrils) can become virtually elastic when Ig domain un/refolding is prevented.     

Walking slope and heavy backpack loads affect torso muscle activity and kinematics

The independent effects of sloped walking or carrying a heavy backpack on posture and torso muscle activations have been reported. While the combined effects of sloped walking and backpack loads are known to be physically demanding, how back and abdominal muscles adapt to walking on slopes with heavy load is unclear. This study quantified three-dimensional pelvis and torso kinematics and muscle activity from longissimus, iliocostalis, rectus abdominis, and external oblique during walking on 0° and ± 10° degree slopes with and without backpack loads using two different backpack configurations (hip-belt assisted and shoulder-borne). Iliocostalis activity was greater during downhill and uphill compared to level walking, but longissimus was only greater during uphill. Rectus abdominis activity was greater during downhill and uphill compared to level, while external oblique activity decreased as slopes progressed from down to up. Longissimus, but not iliocostalis, activity was reduced during both backpack configurations compared to walking with no pack. Hip-belt assisted load carriage required less rectus abdominis activity compared to using shoulder-borne only backpacks; however, external oblique was not influenced by backpack condition. Our results revealed different responses between iliocostalis and longissimus, and between rectus abdominis and external obliques, suggesting different motor control strategies between anatomical planes.     

The functions of the lumbar spine during stepping in the cat

To examine the functional roles played by the lumbar spine during overground stepping, seven adult cats were run in electromyographic (EMG) experiments. Recordings were made bilaterally from mm. iliocostalis, longissimus dorsi and multifidus at a single vertebral level (L₃) and from m. rectus abdominis. Stepping movements were monitored synchronously either by videotape or by high speed cinematography. During alternate use of the hindlimbs (walking and trotting), both epaxial and abdominal muscles were active bilaterally and biphasically. During in-phase use of the hindlimbs (galloping and half-bounding), single bursts of activity were observed. Phasic bursts of activity in rectus abdominus were reciprocal to those of epaxial muscles. Second bursts of activity in either group were noted infrequently. Recordings from the same back muscle at several vertebral levels indicated little difference from these patterns. Movements of the lumbar spine during galloping and half-bounding steps, both angular and linear, are easily correlated with muscle activity patterns. Movements of the lumbar spine during walking and trotting show no particular pattern. Only small angular and linear movements are found. It is concluded that the lumbar spine contributes substantially to step length and limb speed during galloping and half-bounding steps and the epaxial and abdominal musculature may also act as elastic bodies. During walking and trotting steps, the epaxial muscles are proposed to act to stabilize the pelvic girdle to provide a firm base for limb muscles which arise on the pelvis and are synchronously active.     

Homologies of the longissimus, iliocostalis, and hypaxial muscles in the anterior presacral region of extant diapsida

Homologies of muscles of the m. longissimus and m. iliocostalis groups in the dorsal and cervical regions, as well as those of the subvertebral muscles and mm. intercostales externi that continue from the dorsal into the cervical regions, in extant Diapsida are proposed based on detailed dissections and published accounts of lepidosaurs, crocodylians, and birds. The morphology of tendons and innervation patterns suggest that the avian "m. iliocostalis" in the dorsal region include the homologs of both m. longissimus and m. iliocostalis in non-avian diapsids. The conserved nature of the morphology of tendons in palaeognath birds also revealed that the avian mm. intertransversarii in the cervical region consist of muscles of the both m. longissimus and m. iliocostalis groups despite having been treated as a single series of muscles, and thus are not homologous with muscles of the same name in Lepidosauria or Crocodylia. The avian mm. inclusi that lie medial to mm. intertransversarii are homologous with mm. intercostales externi in Lepidosauria and mm. intercostales externi and m. scalenus combined in Crocodylia. Innervation patterns suggest that a muscle ("m. iliocostalis capitis") connecting the atlas rib and occiput in Crocodylia includes contributions from the subvertebral layer and m. cucullaris complex, and possibly m. iliocostalis as well. The present findings may serve as a basis for revising the currently used avian nomenclature so that it will reflect homologies of muscles with their non-avian counterparts.