Myofibrillar M-band proteins in rat skeletal muscles during development

Summary The distribution of three myofibrillar M-band proteins, myomesin, M-protein and the muscle isoform of creatine kinase, was investigated with immunocytochemical techniques in skeletal muscles of embryonic, fetal, newborn and four-week-old rats. Furthermore, muscles of newborn rats were denervated and examined at four weeks of age. In embryos, myomesin was present in all myotome muscle fibres of the somites, whereas M-protein was detected only in a small proportion of the myotome muscle fibres and muscle creatine kinase was not detected at all. In fetal and newborn muscles, all fibres contained all three M-band proteins. At four weeks of age, when fibre types (type 1 or slow twitch fibres and type 2 or fast twitch fibres) were clearly discernable, the pattern was changed. Myomesin and muscle creatine kinase were still observed in all fibres, whereas M-protein was present only in type 2 fibres. On the other hand, in muscle fibres denervated at birth all three M-band proteins were still detected. Our results suggest 1) that during the initial stages of myofibrillogenesis expression and incorporation of myomesin into the M-band precede that of M-protein and muscle creatine kinase; 2) that expression and incorporation of all three M-band proteins during fetal development is nerve independent and non coordinated to the expression of different forms of myosin heavy chains, and 3) that the suppression of M-protein synthesis during postnatal development is nerve dependent and reflects the maturation of slow twitch motor units.     

Effects of growth hormone injection to embryos on growth and myosin heavy chain isoforms in growing turkeys (Meleagris gallopavo)

Effects of embryonic imprinting with growth hormone (GH) on growth and myosin heavy chain (MyHC) isoforms in pectoralis muscle were determined by injecting turkey embryos with ovine growth hormone (oGH) at a dose of 10 μg three times a day. Injections were made on days 20 and 26 (Treatment 1), days 14 and 20 (Treatment 2) or days 14 and 26 (Treatment 3) of incubation. In Treatement 1 poults, plasma GH concentrations were elevated at 3 days posthatch and in Treatment 3 poults, plasma GH concentrations were elevated at 15 days posthatch, as compared to control poults. At 4 weeks of age, in males, body weights, shank length and weights of pectoralis, gastrocnemius and sartorius muscles were increased in Treatment 3, and in females, body weights, shank length and weights of gastrocnemius muscle of female turkeys were increased in Treatment 1. The growth rate of female turkeys from 4 weeks through 16 weeks was increased by Treatment 1. Treatment 1 resulted in a delay in the transition from the embryonic MyHC isoform to the neonatal MyHC isoform and to the adult MyHC isoform. Treatment 3 induced an earlier appearance of the adult MyHC isoform. No effects on body and muscle growth and MyHC isoforms were observed by Treatment 2.     

Myofibrillar M-band proteins in rat skeletal muscles during development.

The distribution of three myofibrillar M-band proteins, myomesin, M-protein and the muscle isoform of creatine kinase, was investigated with immunocytochemical techniques in skeletal muscles of embryonic, fetal, newborn and four-week-old rats. Furthermore, muscles of newborn rats were denervated and examined at four weeks of age. In embryos, myomesin was present in all myotome muscle fibres of the somites, whereas M-protein was detected only in a small proportion of the myotome muscle fibres and muscle creatine kinase was not detected at all. In fetal and newborn muscles, all fibres contained all three M-band proteins. At four weeks of age, when fibre types (type 1 or slow twitch fibres and type 2 or fast twitch fibres) were clearly discernable, the pattern was changed. Myomesin and muscle creatine kinase were still observed in all fibres, whereas M-protein was present only in type 2 fibres. On the other hand, in muscle fibres denervated at birth all three M-band proteins were still detected. Our results suggest 1) that during the initial stages of myofibrillogenesis expression and incorporation of myomesin into the M-band precede that of M-protein and muscle creatine kinase; 2) that expression and incorporation of all three M-band proteins during fetal development is nerve independent and non coordinated to the expression of different forms of myosin heavy chains, and 3) that the suppression of M-protein synthesis during postnatal development is nerve dependent and reflects the maturation of slow twitch motor units.     

Some properties of different skeletal muscle fiber types: comparison of reference bases

Several biochemical components of the white portion of the gastrocnemius (WGM), plantaris (PM), and soleus (SM) muscles of the rat and middle gluteal (MGM) muscle of the horse were compared based on wet and dry weight, protein, and total creatine concentrations ([TCr]). The water content was similar for the rat hindlimb muscles, however, the concentrations of protein, ATP, phosphocreatine (PCr), creatine, and glycogen ranked as SM less than PM less than WGM for all reference bases except total creatine. In contrast, concentrations of ATP, creatine, and PCr were similar in all muscles studied when expressed as ratios of [TCr]. Horse MGM had the lowest percent of water and protein per gram wet or dry weight but highest glycogen concentration of the muscles studied, irrespective of the reference base used to express concentrations. Coefficients of variation were lowest when muscle constituents were related to [TCr]. It is concluded that expressing muscle constituents relative to total creatine results in the smallest variation and is a good method for making comparisons between muscles of similar fiber composition. However, essential information concerning different types of muscle may be lost when this reference base is used.     

The effect of age and castration on the growth rate,blood lipid profile,liver histology and feed conversion in Green-legged Partridge cockerels and capons

The surgical castration of male chickens induces hormonal changes, which permanently influence metabolic processes in birds. The aim of this study was to determine the effect of age and castration on the growth rate, feed conversion, lipid profile and histopathological changes in the livers of cockerels and capons. The experimental materials comprised male chickens of the Green-legged Partridge breed (old traditional Polish chicken breed), raised to 28 weeks of age. At 8 weeks of age, 100 birds were castrated. Caponization had a significant effect on the plasma concentrations of total cholesterol, high-density lipoprotein cholesterol, low-density lipoprotein cholesterol and triacylglycerols (P<0.05). Fatty degeneration and lymphoid cell infiltration were observed more frequently in the livers of capons than cockerels. Capon carcasses were characterized by increased deposition of abdominal and subcutaneous fat (P<0.05). Total meat weight in the carcasses of cockerels and capons was similar, but the proportions of muscles were different. From 20 weeks of age, the weight of breast muscles was higher, and the weight of leg muscles was lower in capons than in cockerels (P<0.05). Capons were characterized by higher liver weight, higher gizzard weight and lower heart weight than cockerels (P<0.05). The feed conversion ratio (kg/kg BW) was similar in intact cockerels and capons. The values of carcass quality parameters and feed conversion ratio as well as histopathological changes in the liver indicate that Green-legged Partridge capons should be slaughtered at 20 to 24 weeks of age.     

Neurochemical effects of extended exposure to white spirit vapour at three concentration levels

Male Wistar rats were exposed to 575 (100 ppm), 2875 (500 ppm) or 5750 mg/m³ (1000 ppm) white spirit vapour for 4–17 weeks 5 days a week, 6 h daily. Perirenal fat solvent concentration corresponded in composition and concentration to those of the vapour at all times. The neurochemical effects included a dose-dependent decrease in the cerebellar succinate dehydrogenase activity for 8 weeks while creatine kinase activity increased after 12 weeks. The specific creatine kinase activity in the glial cell fraction, a marker for astroglia, did not increase suggesting proliferation of astroglial cells in the homogenate. The serum creatine kinase activity originating mainly from striated muscle was below the control range at the two higher concentrations after 12 weeks. Simultaneous analyses for isolated muscle membrane sialic acid and uronic acid residues showed decreased concentrations in proportion to lipid phosphorus or total membrane protein. Thus, the white spirit mixture has neurochemical effects possibly caused by paraffins and the same components may have caused the muscle cell membrane effects. The lowest exposure concentration represents a virtual ‘no effect’ level for rats in the 17-week exposure.     

The effect of thyroid hormones on the aerobic development of locomotor and cardiac muscles in the barnacle goose

The role of thyroid hormones in the development of the locomotor and cardiac muscles of the barnacle goose (Branta leucopsis) was investigated. From 2 weeks of age, goslings were treated with thyroxine, triiodothyronine, or methimazole (a thyroid inhibitor). Birds were killed at 6 weeks (n = 5) or 9 weeks (n = 4) and various locomotor and cardiac muscle masses recorded and tissue samples taken for analysis of citrate synthase activity. The effects of thyroxine and triiodothyronine were not significantly different from each other, except in the case of the iliofibularis at 9 weeks. The mass-specific citrate synthase activity of the iliofibularis, semimembranosus muscles and liver were significantly increased by thyroid hormone treatment. Cardiac muscle showed a significant hypertrophy at 9 weeks of age following treatment with thyroxine. Hypothyroidism induced by treatment with methimazole exhibited the greatest effect on the pectoralis muscle, reducing citrate synthase activity by 33%. Mass-specific citrate synthase activity of the pectoralis and pectoralis mass (% body wt.) were found to be highly correlated (r ² = 0.74) at 6 weeks of age. It is suggested that thyroid hormones may be involved in controlling the tissue-specific timing of the maturation of locomotor and cardiac muscles. Accepted: 16 September 1996     

Progressive loss of mitochondrial creatine phosphokinase activity in muscular dystrophy.

Mitochondria were isolated from the pectoralis and gastrocnemius muscles of chickens with a hereditary muscular dystrophy, and age-matched controls. In the pectoralis, for dystrophic birds aged 0.12, 0.25, 0.55, and 1.55 yr, the creatine phosphokinase activity of the intact mitochondria, expressed in terms of pellet protein, was 69%, 45%, 24%, and 13% as great, respectively, as that of the controls. The corresponding figures for the gastrocnemius were 79%, 46%, 51%, and 28%. The mitochondria from dystrophic muscles exhibited satisfactory respiratory control ratios, P:0 ratios, and state 3 respiratory rates. To check whether their apparent loss of creatine phosphokinase activity was due to the presence of increasing amounts of non-mitochondrial pellet protein, the state 3 respiratory rate was used as a mitochondrial marker; the rates per mg protein were similar in mitochondria from normal and dystrophic muscles of each age group.     

The effect of creatine supplementation on glucose uptake in rat skeletal muscle

Glucose transport in muscle is a function of the muscle metabolic state, as evidenced by the increase in glucose transport which occurs with conditions of altered aerobic metabolism such as hypoxia or contractile activity. The energy state of the muscle can be determined by the muscle phosphocreatine concentration. Dietary supplementation of creatine has been shown to increase both phosphocreatine (PCr) and creatine (TCr) levels in muscle, although not in the same proportion, so that the PCr/TCr ratio falls suggesting an altered energy state in the cell. The purpose of this study was to determine the effect of increased creatine content on glucose uptake in muscle. PCr and TCr were determined in plantaris muscles from rats following five weeks of dietary supplementation of creatine monohydrate (300 mg/kg/day). (3)H-2-deoxyglucose uptake was measured in epitrochlearis muscles incubated in the presence or absence of a maximally stimulating dose of insulin. Despite a significant increase in creatine content in muscle, neither basal nor insulin-stimulated glucose uptake was altered in creatine supplemented rats. Since PCr levels were not increased with creatine supplementation, these results suggest that the actual concentration of PCr is a more important determinant of glucose uptake than the PCr/TCr ratio.     

Structure of skeletal muscles in leghorn type chicken from conservative and parent flocks

Intensive selection conducted within closed populations has led to the creation of specialized chicken strains that differ significantly in meat yield and reproduction performance. The effect of the selection conducted on the birds is differentiation identified not only on the molecular but also on the cellular level, among other things in the skeletal muscles. The aim of this study was to compare the structure of chosen homological skeletal muscles from Leghorn chickens (LSL), originating from parent flock, intensively selected for reproductive traits and from conservative flock (G99), unselected for many generations. The structure of musculus pectoralis superficialis and musculus biceps femoris (the thickness of the muscle fibres and the share of the fibre types in the bundle) in 8 and 20 week old chickens was compared. A significant impact of the origin on all examined slaughter parameters was recorded. Body weight before slaughter, carcass weight and the weight of breast and leg muscles in 8 weeks old LSL chicken made up from 60% to about 85% of the respective values in the G99 Leghorn. Lack of red fibres in the breast muscles of all the individuals from the parental flock (LSL) was noted, whereas in 12 individuals (among 24) from the conservative flock (G99), red fibres were observed in this muscle from 2.75% up to 7.09%. White fibres in 8 week old chicks were always thicker, both in pectoralis superficialis and biceps femoris muscle in birds with higher body weight as well as higher weight of breast and legs muscles, i.e. in chicks from conservative flock (G99), P<0.01. However, in 20 week old birds, the diameters of the white fibres were similar in both groups. Also the diameters of the red fibres in musculus biceps femoris in 8 week old chickens were higher in cockerels and pullets from conservative flock (G99).     

Ontogeny of catabolic and morphological properties of skeletal muscle of the red-winged blackbird (Agelaius phoeniceus)

Thermogenic capabilities of red-winged blackbirds improve markedly during their 10-12-day nestling period, especially between day 5 and day 8. The time course of improvements may be determined by the maturation of skeletal muscles involved in shivering thermogenesis, particularly the pectoralis muscles. To test this hypothesis, morphological and biochemical changes in pectoral and leg muscles were measured in young and adult blackbirds. Both muscles grew disproportionately relative to body mass. The pectoralis consisted entirely of fast-twitch fibers, predominantly fast oxidative glycolytic. In contrast, the gastrocnemius muscle consisted of a mixture of slow and fast fibers (predominantly fast glycolytic). Although fiber composition was constant, both cross-sectional area and density of fibers increased with age in both muscles. Catabolic capacities of the pectoralis increased significantly (approximately 7-8-fold) throughout the nestling period, most abruptly after day 3 (citrate synthase, CS) or day 4 (3-hydroxacyl-CoA-dehydrogenase, HOAD). Myofibrillar ATPase activities in the pectoralis were initially low, but increased after day 5. Further increases in CS and myofibrillar ATPase activities occurred in the pectoralis after fledging. CS and HOAD activities in the leg were much lower, but myofibrillar ATPase activities were remarkably similar in the two muscles, differing only in adults. These results are consistent with the hypothesis that the development of endothermy is dependent on the morphological and biochemical maturation of skeletal muscles important in thermogenesis.     

High beta-hydroxybutyrate concentration in liver and skeletal muscle of newly hatched chicks

Characteristic changes in ketone body concentrations in blood, liver, and skeletal muscle were investigated in detail in newly hatched chicks. The concentration of beta-hydroxybutyrate in the blood was maximal at hatch (0 day), markedly decreased to 3 days, then maintained at low levels, up to 14 days of age. The concentration of acetoacetate in blood, on the other hand, did not change after hatching but remained lower than that of beta-hydroxybutyrate at all ages. In liver and muscles, the concentration of beta-hydroxybutyrate changed in a manner similar to that in the blood. The muscle to blood ratio of the beta-hydroxybutyrate concentration on days -1 and 0 was significantly higher than those at 1 through 14 days post-hatch. These results show that newly hatched chicks have the same high ketone body concentrations in the skeletal muscle, blood and liver. It is, hence, suggested that uptake of beta-hydroxybutyrate by muscles is substantial or that ketogenesis, if any, occurs in muscles immediately before and after hatching of chicks.     

Comparison of muscle activation using various hand positions during the push-up exercise

Popular fitness literature suggests that varied hand placements during push-ups may isolate different muscles. Scientific literature, however, offers scant evidence that varied hand placements elicit different muscle responses. This study examined whether different levels of electromyographic (EMG) activity in the pectoralis major and triceps brachii muscles are required to perform push-ups from each of 3 different hand positions: shoulder width base, wide base, and narrow base hand placements. Forty subjects, 11 men and 29 women, performed 1 repetition of each push-up. The EMG activity for subjects' dominant arm pectoralis major and triceps brachii was recorded using surface electrodes. The EMG activity was greater in both muscle groups during push-ups performed from the narrow base hand position compared with the wide base position (p < 0.05). This study suggests that, if a goal is to induce greater muscle activation during exercise, then push-ups should be performed with hands in a narrow base position compared with a wide base position.     

No effects of lifelong creatine supplementation on sarcopenia in senescence-accelerated mice (SAMP8)

Oral creatine supplementation can acutely ameliorate skeletal muscle function in older humans, but its value in the prevention of sarcopenia remains unknown. We evaluated the effects of lifelong creatine supplementation on muscle mass and morphology, contractility, and metabolic properties in a mouse model of muscle senescence. Male senescence-accelerated mice (SAMP8) were fed control or creatine-supplemented (2% of food intake) diet from the age of 10 to 60 wk. Soleus and extensor digitorum longus muscles were tested for in vitro contractile properties, creatine content, and morphology at weeks 25 and 60. Both muscle types showed reduced phosphocreatine content at week 60 that could not be prevented by creatine. Accordingly, age-associated decline in muscle mass and contractility was not influenced by treatment. Aged soleus muscles had fewer and smaller fast-twitch glycolytic fibers irrespective of treatment received. It is concluded that lifelong creatine supplementation is no effective strategy to prevent sarcopenia in senescence-accelerated mice.     

31P NMR study of postmortem metabolism in porcine and bovine muscles

31P NMR spectroscopy was used to evaluate interspecies differences in muscle fibre types and related postmortem metabolism. M. longissimus thoracis (MLT) and m. pectoralis superficialis (MPS) of bulls and MLT of pigs were investigated. In perchloric acid extracts NMR resonances for sugar phosphates (SP), inorganic phosphate (Pi), glycerophosphorylcholine (GPC), phosphocreatine (PCr), adenosine triposphate (ATP), adenosine diphosphate (ADP) as well as for NAD+/NADH could be distinguished. Also, glycogen and lactate contents and pH were determined. The relative contents of phosphorus compounds in bovine muscles of similar participation of muscle fibre are similar. Bovine muscles contain a relatively large proportion of PCr (48% of all phosphates 15 minutes post-mortem in MPS) whereas porcine MLT show lower PCr content (11% 15 minutes post-mortem). On the other hand, the ATP content is relatively higher in porcine MLT when compared with bovine muscles in the early phases of the postmortem processes. No NMR-detectable levels of GPC were measured in porcine MLT in contrast to bovine muscles. This suggests that the GPC content does not depend solely on the fibre participation but is also animal species determined. The 24 hour postmortem metabolism patterns of bovine and porcine muscles have many common traits. CP disappeared first followed by ATP. Simultaneously, the Pi concentrations increased. However, the content of SP remained relatively constant in porcine, but not in bovine muscles where it increased only gradually. The significantly higher concentrations of SP and lactate as well as the lower values of glycogen and pH measured for porcine as compared with bovine muscles suggest an enhanced glycolysis during the early phases of postmortem processes in porcine muscles.     

The morphological development of the locomotor and cardiac muscles of the migratory barnacle goose (Branta Leucopsis)

The masses of the locomotor and acardiac muscles of wild barncale goose gollings, from migratory population, were examined systematically during development and their values compared to those of pre-migratory geese. Pre-flight development was typified by approximately linear increases of body, leg, and heart ventricular mass with respect to age. Flight muscle showed an exponential increase in mass. Pectoralis muscle mass was 14.2 ± 0.3% of body mass (1297 ± 73g, n=7) in early flying goslings compared to 16.6 ± 0.3% of body mass (2318 ± 109g, n=8) in pre-migratory geese. Post-flight development was typified by stasis of leg muscle mass but hypertrophy of Ventricular and pectoralis muscle mass in proportion to body mass. Ventricular mass relative to body mass showed the lowest values at 5 weeks of age (0.62 ± 0.01%) with peak values at 1 week of age (10.4 ± 0.04%). The latter may be associated with both requirements of thermoregualation in these precocial, arctic breeding geese and the need to forage approximately 24 hours post-hatch. Peak values for leg muscle mass, relative to body mass, were found at 3 weeks of age (12.7 ± 0.36%), with lowest values in the pre-migratory geese (6.7 ± 0.21%), while peak values for pectoralis muscle mass were expressed in the premigratory geese with lowest values at 1 week of age (0.94 ± 0.07%). Ventricular mass was proportional to leg muscle mass up to 5 weeks of age (M_v= 0.38M_t^0.68, r²=0.95), but subsequent increase in ventricular mass was proportional to pectoralis muscle mass (M_v= 0.25M_p^0.73, r²= 0.81).     

Thyroidal involvement in the expression of avian muscular dystrophy

We showed previously that propylthiouracil (PTU), a thyroid inhibitor, could alleviate several major signs of hereditary muscular dystrophy in chickens. The goals of the present investigation were to: (1) determine whether a nearly athyroid condition (achieved within two days after hatching by surgical thyroidectomy plus PTU) during an 11-day period beneficially affects the dystrophic condition when followed by triiodothyronine (T3) replacement to 33 days of age; (2) determine the beneficial effects on the expression of avian dystrophy when the thyroidectomized-PTU-treated chickens received a wide range of moderate to low T3 replacement doses beginning by two days after thyroidectomy; and (3) examine the thyroid hormone receptor system in dystrophic muscle for a possible abnormality. Thyroid deprivation increased muscle function (righting ability) and reduced plasma creatine kinase activity in dystrophic chickens. The major thyroid-related abnormality in dystrophic pectoralis muscles was an increased maximum binding capacity of solubilized nuclear T3 receptors.     

Changes of tissue creatine concentrations upon oral supplementation of creatine-monohydrate in various animal species

Creatine is a nutritional supplement with major application as ergogenic and neuroprotective substrate. Varying supplementation protocols differing in dosage and duration have been applied but systematic studies of total creatine (creatine and phosphocreatine) content in the various organs of interest are lacking. We investigated changes of total creatine concentrations in brain, muscle, heart, kidney, liver, lung and venous/portal plasma of guinea pigs, mice and rats in response to 2-8 weeks oral creatine-monohydrate supplementation (1.3-2 g/kg/d; 1.4-2.8% of dietary intake). Analysis of creatine and phosphocreatine content was performed by high performance liquid chromatography. Total creatine was determined as the sum of creatine and phosphocreatine. Presupplementation total creatine concentrations were high in brain, skeletal and heart muscle (10-22 micromol/g wet weight), and low in liver, kidney and lung (5-8 micromol/g wet weight). During creatine supplementation, the relative increase of total creatine was low (15-55% of presupplementation values) in organs with high presupplementation concentrations, and high (260-500% of presupplementation values) in organs with low presupplementation concentrations. The increase of total creatine concentrations was most pronounced after 4 weeks of supplementation. In muscle, brain, kidney and lungs, an additional increase (p<0.01) was observed between 2-4 and 2-8 weeks of supplementation. Absolute concentrations of phosphocreatine increased, but there was no increase of the relative (percentual) proportion of phosphocreatine (14-45%) during supplementation. Statistical comparison of total creatine concentrations across the species revealed no systematically differences in organ distribution and in time points of supplementation. Results suggest that in organs with low presupplementation creatine levels (liver, kidney), a major determinant of creatine uptake is an extra-intracellular concentration gradient. In organs with high presupplementation total creatine levels like brain, skeletal and heart muscle, the maximum capacity of creatine accumulation is low compared to other organs. A supplementation period of 2 to 4 weeks is necessary for significant augmentation of the creatine pool in these organs.     

Postnatal development of thiamine metabolism in rat skeletal muscle.

1. The activities of 2-oxoglutarate dehydrogenase, transketolase, thiamine pyrophosphokinase and thiamine triphosphatase and the concentrations of thiamine phosphates were almost the same between rat extensor digitorum longus and soleus muscles at 2 weeks of age. 2. These enzyme activities changed after 3 weeks of age in a different way depending on the muscle phenotype. 3. Thiamine diphosphate level and the activity of 2-oxoglutarate dehydrogenase increased only in soleus muscle and thiamine triphosphate level increased only in extensor digitorum longus during development.     

Electromyography of pongid shoulder muscles III. Quadrupedal positional behavior

Electromyographic (EMG) recordings were taken from 14 shoulder muscles (or major parts of them) in a gorilla, a chimpanzee and an orangutan as they stood quadrupedally and tripedally, descended from elevated substrates, crutch-walked, and progressed quadrupedally on inclined and level substrates. In the African apes, low potentials commonly (but not always) occurred in the sternocostal pectoralis major, anterior deltoid, supraspinatus and subscapularis muscles during quadrupedal stance. The quadrupedal orangutan always exhibited low potentials in the pectoralis major muscle and EMG activity commonly occurred in her supraspinatus and subscapularis muscles. Quiescent tripedal stances were not accompanied by striking changes in EMG patterns from those which characterized quadrupedal stances. Per contra, eccentric loadings of the forelimb during descents from elevated substrates generally recruited notable EMG activity in the deltoid, supraspinatus and, to a lesser extent, infraspinatus muscles of the three pongid apes. The pectoralis major and caudal serratus anterior muscles were much more active in Pongo and Pan during these descents. Supportive segments of quadrupedal locomotive cycles were generally accompanied by EMG activity in the pectoralis major, intermediate and posterior deltoid and supraspinatus muscles. The intermediate and posterior deltoid muscles were characteristically active during pre-release of the hand and early swing phase. The cranial trapezius and supraspinatus muscles also may act during early swing phase. We conclude that the pectoralis major and perhaps the supraspinatus and subscapularis might serve regularly as postural muscles during static terrestrial quadrupedalism in pongid apes. The lack of dramatic differences between the EMG patterns exhibited during fist-walking versus knuckle-walking indicates that an evolutionary transformation from a shoulder complex like that of Pongo to ones like Pan or vice versa need not entail major changes in myological features.