Neuromuscular organization of avian flight muscle: architecture of single muscle fibres in muscle units of the pectoralis (pars thoracicus) of pigeon (Columba livia)

The M. pectoralis (pars thoracicus) of pigeons (Columba livia) is comprised of short muscle fibres that do not extend from muscle origin to insertion but overlap ''in-series''. Individual pectoralis motor units are limited in territory to a portion of muscle length and are comprised of either fast twitch, oxidative and glycolytic fibres (FOG) or fast twitch and glycolytic fibres (FG). FOG fibres make up 88 to 90% of the total muscle population and have a mean diameter one-half of that of the relatively large FG fibres. Here we report on the organization of individual fibres identified in six muscle units depleted of glycogen, three comprised of FOG fibres and three comprised of FG fibres. For each motor unit, fibre counts revealed unequal numbers of depleted fibres in different unit cross-sections. We traced individual fibres in one unit comprised of FOG fibres and a second comprised of FG fibres. Six fibres from a FOG unit (total length 15.45 mm) ranged from 10.11 to 11.82 mm in length and averaged (± s.d.) 10.74 ± 0.79 mm. All originated bluntly (en mass) from a fascicle near the proximal end of the muscle unit and all terminated intramuscularly. Five of these ended in a taper and one ended bluntly. Fibres coursed on average for 70% of the muscle unit length. Six fibres from a FG unit (total length 34.76 mm) ranged from 8.97 to 18.38 mm in length and averaged 15.32 ± 3.75 mm. All originated bluntly and terminated intramuscularly; one of these ended in a taper and five ended bluntly. Fibres coursed on average for 44% of the muscle unit length. Because fibres of individual muscle units do not extend the whole muscle unit territory, the effective cross-sectional area changes along the motor unit length. These non-uniformities in the distribution of fibres within a muscle unit emphasize that the functional interactions within and between motor units are complex.     

Lack of fiber type selectivity during reinnervation of neonatal rabbit soleus muscle

Fast and slow contracting fibers in neonatal mammalian skeletal muscle are each innervated in a highly specific manner by motor neurons of the corresponding type, even at an age when polyinnervation is widespread. Chemospecific recognition is a possible mechanism by which this pattern of innervation could be established. We have investigated this possibility by studying the degree of specificity during reinnervation of rabbit soleus muscle following nerve crush on Postnatal Day 1 or 4. We assayed fiber type composition by measuring the twitch rise times of motor units within 2 days of the onset of functional reinnervation (5-6 days after nerve crush). In contrast to the broad, bimodal distribution of single motor unit twitch rise times seen in normal muscles, motor units in reinnervated muscles yielded a narrower, unimodal distribution of rise times. Rise times of reinnervated units were intermediate to those of normal fast and slow units, suggesting that reinnervated units were composed of a mixture of fast and slow contracting fibers. An alternative possibility, that specific reinnervation was masked by contractile dedifferentiation of muscle fibers, was examined by maintaining a transmission blockade induced by botulinum toxin poisoning for an equivalent interval. Twitch rise times of treated motor units exhibited the distinctly bimodal distribution characteristic of normal muscles, suggesting that muscle fibers can retain contractile diversity during a transient period of denervation. We carried out computer simulations to estimate the amount of rise time diversity induced by varying degrees of specificity during reinnervation. Based on this analysis, we conclude that there is little if any selective reinnervation of muscle fiber types at the ages studied.     

Motor unit composition has little effect on the short-range stiffness of feline medial gastrocnemius muscle.

Studies on skinned fibers and single motor units have indicated that slow-twitch fibers are stiffer than fast-twitch fibers. This suggests that skeletal muscles with different motor unit compositions may have different short-range stiffness (SRS) properties. Furthermore, the natural recruitment of slow before fast motor units may result in an SRS-force profile that is different from electrical stimulation. However, muscle architecture and the mechanical properties of surrounding tissues also contribute to the net SRS of a muscle, and it remains unclear how these structural features each contribute to the SRS of a muscle. In this study, the SRS-force characteristics of cat medial gastrocnemius muscle were measured during natural activation using the crossed-extension reflex, which activates slow before fast motor units, and during electrical activation, in which all motor units are activated synchronously. Short, rapid, isovelocity stretches were applied using a linear puller to measure SRS across the range of muscle forces. Data were collected from eight animals. Although there was a trend toward greater stiffness during natural activation, this trend was small and not statistically significant across the population of animals tested. A simple model, in which the slow-twitch fibers were assumed to be 30% stiffer than the fast-twitch fibers, was used to simulate the experimental results. Experimental and simulated results show that motor unit composition or firing rate has little effect on the SRS property of the cat MG muscle, suggesting that architectural features may be the primary determinant of SRS.     

Ultrastructural diversity in motor units of crustacean stomach muscles

The physiological and ultrastructural properties of muscle fiber.s comprising three motor units in the gastric mill of blue crabs are described. In their contractile properties muscle fibers in all motor units are similar and resemble the slow type fibers in crustacean limb muscles. The majority of fibers generate large excitatory post-synaptic potentials which do not facilitate strongly. Structurally two types of fibers are found. The one type has long sarcomeres (greater than 6 mum), thin to thick myofilament ratios of 5-6:1 and diads located near the ends of the A-band. The other type has shorter sarcomeres (less than 6 mum), thin to thick myofilament ratios of 3:1 and diads located at mid sarcomere level. Both types of fibers occur within a single motor unit and this differs from the vertebrate situation. Furthermore, the finding of fibers with a low thin to thick myofilament ratio of 3:1 demonstrates that they are not exclusive to fast type crustacean muscle but also occur in slow stomach muscles.     

Functional and clinical significance of the architecture of human skeletal muscles

The architectural properties of the triceps surae muscle were studied in vivo in groups of healthy subjects (eight men) and patients with locomotor function disorders (four men and four women) with the ankle joint positioned at a plantar flexion 0° and the knee set at 90° (neutral position). In this position, using ultrasonic scanning, longitudinal ultrasonic images of the medial gastrocnemius (MG), lateral gastrocnemius (LG), and soleus (Sol) muscles were obtained when the subject was relaxed (the passive state) or performed isometric plantar flexion (50% of the maximum voluntary contraction (MVC), the active state). The fascicle lengths, fascicle angles, and muscle thickness were determined. In the passive state, the fascicle lengths of the MG, LG, and Sol muscles in the group of healthy subjects were 33, 35, and 30 mm and the pennation angle, 25°, 19°, and 25°; in the group of patients with motor disorders, 38, 39, and 29 mm and 21°, 19°, and 24°, respectively. The MG, LG, and Sol thicknesses in the group of healthy subjects were 15, 13, and 12 mm, and in the group of patients with motor disorders, 14, 12, and 14 mm, respectively. In the active state (50% of the MVC), the MG, LG, and Sol fiber lengths in the group of healthy subjects shortened by 31, 24, and 18%; the fiber pennation angle increased by 60, 41, and 41%, respectively. In the group of patients with motor disorders, the fiber lengths shortened by 28, 14, and 18% and the fiber pennation angle decreased by 28, 26, and 36%, respectively. The MG, LG, and Sol thicknesses in the group of healthy subjects increased by 9, 22, and 18%, while in the group of patients with motor disorders the thickness decreased by 4% in the MG and increased by 11 and 4% in the LG and Sol muscles, respectively. Different fiber lengths and pennation angles and their changes upon contraction might be related to differences in the force-producing capabilities of the muscles and the viscoelastic properties of muscle tendons and aponeuroses.     

Asymmetric deformation of contracting human gastrocnemius muscle

Muscle fiber deformation is related to its cellular structure, as well as its architectural arrangement within the musculoskeletal system. While playing an important role in aponeurosis displacement, and efficiency of force transmission to the tendon, such deformation also provides important clues about the underlying mechanical structure of the muscle. We hypothesized that muscle fiber cross section would deform asymmetrically to satisfy the observed constant volume of muscle during a contraction. Velocity-encoded, phase-contrast, and morphological magnetic resonance imaging techniques were used to measure changes in fascicle length, pinnation angle, and aponeurosis separation of the human gastrocnemius muscle during passive and active eccentric ankle joint movements. These parameters were then used to subsequently calculate the in-plane muscle area subtended by the two aponeuroses and fascicles and to calculate the in-plane (dividing area by fascicle length), and through-plane (dividing muscle volume by area) thicknesses. Constant-volume considerations of the whole-muscle geometry require that, as fascicle length increases, the muscle fiber cross-sectional area must decrease in proportion to the length change. Our empirical findings confirm the definition of a constant-volume rule that dictates that changes in the dimension perpendicular to the plane, i.e., through-plane thickness, (-6.0% for passive, -3.3% for eccentric) equate to the reciprocal of the changes in area (6.8% for passive, 3.7% for eccentric) for both exercise paradigms. The asymmetry in fascicle cross-section deformation for both passive and active muscle fibers is established in this study with a ～22% in-plane and ～6% through-plane fascicle thickness change. These fiber deformations have functional relevance, not only because they affect the force production of the muscle itself, but also because they affect the characteristics of adjacent muscles by deflecting their line of pull.     

The histochemical profile of the rat extensor digitorum longus muscle differentiates after birth and dedifferentiates in senescence

Age dependent motor unit dedifferentiation is a key component of impaired muscle function in advanced age. Here, we tested the hypothesis that rat muscle histochemical profile during the lifespan of an individual has an age-specific pattern since comprehensive longitudinal studies of muscle differentiation after birth and dedifferentiation in advanced age are scarce. Our results show that extensor digitorum longus muscle (EDL) is comprised only of two fiber types after birth, type slow-oxidative (SO) and type SDH-intermediate (SDH-INT), the latter being indicative for the presence of polyneuronal innervation. In contrast to the constantly growing cross-sectional area of the muscle fibers, a dramatic decrease in SDH-INT proportion occurs between day 14 and 21 after birth resulting in a complete loss of fiber type SDH-INT at the age of 90 days (p<0.05). At the age of 270 days, the fiber type composition of rat EDL dedifferentiates as shown by the reappearance of the SDH-INT type with a further increase at the age of 540 days (p<0.05). These changes in histochemical fiber type spectra are brought about by fiber type conversion within the fast twich fibers. The findings of the present study provide further evidence that fiber type conversion is a basic mechanism leading to motor unit differentiation and dedifferentiation during ontogenesis. Fiber type conversion shows a distinct time specific pattern and is also characteristic for motor unit regeneration after peripheral nerve repair. Factors that influence fiber type conversion and thereby motor unit organization may provide a future therapeutic option to enhance the regenerative capacity of motor units.     

Simulation of motor unit recruitment and microvascular unit perfusion: spatial considerations

Fuglevand, Andrew J., and Steven S. Segal. Simulationof motor unit recruitment and microvascular unit perfusion: spatial considerations. J. Appl. Physiol.83(4): 1223-1234, 1997.Muscle fiber activity is the principalstimulus for increasing capillary perfusion during exercise. Thecontrol elements of perfusion, i.e., microvascular units (MVUs), supplyclusters of muscle fibers, whereas the control elements of contraction,i.e., motor units, are composed of fibers widely scattered throughoutmuscle. The purpose of this study was to examine how the discordantspatial domains of MVUs and motor units could influence the proportion of open capillaries (designated as perfusion) throughout a muscle crosssection. A computer model simulated the locations of perfused MVUs inresponse to the activation of up to 100 motor units in a muscle with40,000 fibers and a cross-sectional area of 100 mm². The simulation increasedcontraction intensity by progressive recruitment of motor units. Foreach step of motor unit recruitment, the percentage of active fibersand the number of perfused MVUs were determined for several conditions:1) motor unit fibers widely dispersed and motor unit territories randomly located (whichapproximates healthy human muscle),2) regionalized motor unitterritories, 3) reversed recruitmentorder of motor units, 4) denselyclustered motor unit fibers, and 5)increased size but decreased number of motor units. The simulationsindicated that the widespread dispersion of motor unit fibersfacilitates complete capillary (MVU) perfusion of muscle at low levelsof activity. The efficacy by which muscle fiber activity inducedperfusion was reduced 7- to 14-fold under conditions that decreased thedispersion of active fibers, increased the size of motor units, orreversed the sequence of motor unit recruitment. Such conditions aresimilar to those that arise in neuromuscular disorders, with aging, orduring electrical stimulation of muscle, respectively.

     

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

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

AXIAL MUSCULATURE IN THE DOLPHIN (TURSIOPS TRUNCATUS): SOME ARCHITECTURAL AND HISTOCHEMICAL CHARACTERISTICS

In view of the supposition that a dolphin can swim faster than would be predicted based on its physical features and presumed muscle power potential, studies were initiated to reevaluate the assumptions made in reaching these conclusions. Several previous studies have shown that the architectural and histochemical properties of a skeletal muscle dictate its force, velocity and displacement properties. This study examined the muscle fiber lengths and tendon arrangements of the dorsal and ventral axial muscles in dolphins ( Tursiops truncatus ). Fiber type and fiber size distributions were determined to reflect the general biochemical characteristics of the musculature. The dorsal muscles had a higher mean fiber length (167 Vs. 90 mm) and the range within and across different dorsal muscles was less (141–199 vs. 37–185 mm) than in the ventral muscles. Both the dorsal and ventral muscles consisted of an overall mean of 50 percent slow twitch and 50 percent fast twitch fiber types. The fast twitch fibers were 67 percent larger (2,200 vs. 1,317 μ m ²) than the slow twitch fibers in the ventral and 38 percent larger (1,213 Vs. 879 μm²) in the dorsal muscles. In addition, the mean cross sectional area of the fibers in the ventral muscles was approximately 65 percent greater (1,750 vs. 1,072 μm²) than those in the dorsal muscles. The shorter, larger-diameter fibers of the ventral musculature give it a greater potential for force production for a given amount of muscle mass. In contrast, the dorsal muscles appear to be designed to optimize velocity and displacement, ( i.e. , longer fibers). These findings contribute to the information necessary for the determination of the power potential of the musculature of the dolphin.     

Myosin isozymes in normal and cross-reinnervated cat skeletal muscle fibers

Immunocytochemical characteristics of myosin have been demonstrated directly in normal and cross-reinnervated skeletal muscle fibers whose physiological properties have been defined. Fibers belonging to individual motor units were identified by the glycogen-depletion method, which permits correlation of cytochemical and physiological data on the same fibers. The normal flexor digitorum longus (FDL) of the cat is composed primarily of fast-twitch motor units having muscle fibers with high myosin ATPase activity. These fibers reacted with antibodies specific for the two light chains characteristic of fast myosin, but not with antibodies against slow myosin. Two categories of fast fibers, corresponding to two physiological motor unit types (FF and FR), differed in their immunochemical response, from which it can be concluded that their myosins are distinctive. The soleus (SOL) consists almost entirely of slow-twitch motor units having muscle fibers with low myosin ATPase activity. These fibers reacted with antibodies against slow myosin, but not with antibodies specific for fast myosin. When the FDL muscle was cross-reinnervated by the SOL nerve, twitch contraction times were slowed about twofold, and motor units resembled SOL units in a number of physiological properties. The corresponding muscle fibers had low ATPase activity, and they reacted with antibodies against slow myosin only. The myosin of individual cross-reinnervated FDL muscle units was therefore transformed, apparently completely, to a slow type. In contrast, cross-reinnervation of the SOL muscle by FDL motoneurons did not effect a complete converse transformation. Although cross-reinnervated SOL motor units had faster than normal twitch contraction times (about twofold), other physiological properties characteristic of type S motor units were unchanged. Despite the change in contraction times, cross-reinnervated SOL muscle fibers exhibited no change in ATPase activity. They also continued to react with antibodies against slow myosin, but in contrast to the normal SOL, they now showed a positive response to an antibody specific for one of the light chains of fast myosin. The myosins of both fast and slow muscles were thus converted by cross-reinnervation, but in the SOL, the newly synthesized myosin was not equivalent to that normally present in either the FDL or SOL. This suggests that, in the SOL, alteration of the nerve supply and the associated dynamic activity pattern are not sufficient to completely respecify the type of myosin expressed.     

Three-dimensional analysis of the arrangement and length distribution of fascicles in the triceps muscle of Galea musteloides (Rodentia, Cavimorpha)

Non-uniformity of fascicle parameters (fascicle lengths and orientation) within one skeletal muscle is well known. These parameters have an effect on the physiological cross-sectional area and lengthening rate of the skeletal muscle. Using a binocular microscope with a table driver (q- and p-axes) and vertical drive (v-axis) as a tool for reconstruction of the spatial orientation of single muscle fascicles, we developed an approach for three-dimensional analysis of the arrangement and length distribution in the skeletal muscle of small mammals. Two subunits of the triceps brachii muscle of the Galea musteloides forelimb, triceps longum and triceps laterale, were quantified and compared. Our data show that in the triceps laterale the fascicles are significantly longer (10.23 mm, SD=1.19, n=41) than those in the triceps longum (6.58 mm, SD=2.88, n=39). In the triceps laterale, the fascicle orientation is more or less uniform, whereas, in the triceps longum, there are two areas with different orientation of fascicles: anterior and posterior ones. Different inner architecture of the subunits can be interpreted as an adaptation to the main locomotory function of the triceps muscle, namely production of propulsive force during limb transfer phase and keeping dynamic stability during stance phase. Comparison of our data on the fascicle length and geometry with our previous histochemical results on G. musteloides, shows that the anterior region of the triceps longum, which differs in the fascicle orientation, also contains a significantly larger percent of slow muscle fibres. It is hypothesised here that this small region is involved in keeping posture. Accepted: 16 May 2000     

A micromechanical muscle model for determining the impact of motor unit fiber clustering on force transmission in aging skeletal muscle

This study used a micromechanical finite element muscle model to investigate the effects of the redistribution of spatial activation patterns in young and old muscle. The geometry consisted of a bundle of 19 active muscle fibers encased in endomysium sheets, surrounded by passive tissue to model a fascicle. Force was induced by activating combinations of the 19 active muscle fibers. The spacial clustering of muscle fibers modeled in this study showed unbalanced strains suggesting tissue damage at higher strain levels may occur during higher levels of activation and/or during dynamic conditions. These patterns of motor unit remodeling are one of the consequences of motor unit loss and reinnervation associated with aging. The results did not reveal evident quantitative changes in force transmission between old and young adults, but the patterns of stress and strain distribution were affected, suggesting an uneven distribution of the forces may occur within the fascicle that could provide a mechanism for muscle injury in older muscle.

     

Structure and Function in Vertebrate Skeletal Muscle

SYNOPSIS. The functional diversity of vertebrate skeletal musclelargely depends upon its structure. An important aspect of thisis its hierarchical design. At the cellular level, muscle fibersform three categories whose functional properties grade intoeach other: slow-oxidative fibers with high endurance to fatigue,fast-oxidative\glycolytic fibers also endurant but with greatermetabolic diversity, and fast-glycolytic fibers with limitedendurance but quick response. This partitioning of functionalproperties found among single muscle fibers is conserved ata second level of the structural hierarchy, since the groupof myofibers innervated by a single motor neuron (together calleda motor unit) is composed of the same fiber type. Differentmotor units may be recruited in an orderly pattern dependingupon the functional demands of a particular behavior. Finally,groups of motor units innervated by axons travelling togetherin the primary nerve branches may form discrete neuromuscularcompartments at a third level of structural hierarchy. Differentmotor units may be found in regional arrays in these compartmentsso that slow or fast units tend to be clumped together and maybe recruited together as larger functional units. This hierarchicalorganization of skeletal muscle may be a fundamental vertebrateplan that allows the diversity of functions so evident in vertebratebehavior.     

Musculotendon parameters and musculoskeletal pathways within the human foot

In order to create a flexible model of the foot for dynamic musculoskeletal models, anthropometric data combined with geometric information describing the intrinsic musculature are needed. In this study, the left feet of two male and two female cadavers were dissected to expose the intrinsic musculotendon pathways. Three-dimensional coordinates of bony landmarks, tendon origins, insertions, and via points were digitized to submillimeter accuracy. Muscle architectural parameters were also measured, including volume, weight, and pennation angle and sarcomere, fascicle, and free tendon lengths. Optimal muscle fascicle lengths, pen-nation angles at optimal length, physiological cross-sectional areas (PCSA), and tendon slack lengths were calculated from the directly measured values. Fascicle length and pennation angle varied greatly within each subject. Average fascicle lengths normalized by optimal fascicle length varied between 0.73 and 1.25, with 75% of the formalin-preserved muscles being found in a shortened state. The muscle volume and PCSA also had a large variability within subjects but less variation between subjects. The ratio of tendon slack length to optimal fascicle length was found to vary between 1.05 and 9.56. Using this data, a deformable model of the foot can now be created. It is envisioned that deformable feet will significantly improve stability and realism in models of gait, posture, and sporting activities.     

Muscle fiber and tendon length changes in the human vastus lateralis during slow pedaling.

Muscle fascicle lengths of vastus lateralis (VL) muscle were measured in five healthy men during slow pedaling to investigate the interaction between muscle fibers and tendon. Subjects cycled at a pedaling rate of 40 rpm (98 W). During exercise, fascicle lengths changed from 91 +/- 7 (SE) to 127 +/- 5 mm. It was suggested that fascicles were on the descending limb of their force-length relationship. The average shortening velocity of fascicle was greater than that of muscle-tendon complex in the first half of the knee extension phase and was less in the second half. The maximum shortening velocity of fascicle in the knee extension phase was less than that of muscle-tendon complex by 22 +/- 9%. These discrepancies in velocities were mainly caused by the elongation of the tendinous tissue. It was suggested that the elasticity of VL tendinous tissue enabled VL fascicles to develop force at closer length to their optimal length and kept the maximum shortening velocity of VL fascicles low during slow pedaling.     

Detection of motor unit action potentials with surface electrodes: influence of electrode size and spacing

A model of the motor unit action potential was developed to investigate the amplitude and frequency spectrum contributions of motor units, located at various depths within muscle, to the surface detected electromyographic (EMG) signal. A dipole representation of the transmembrane current in a three-dimensional muscle volume was used to estimate detected individual muscle fiber action potentials. The effects of anisotropic muscle conductance, innervation zone location, propagation velocity, fiber length, electrode area, and electrode configuration were included in the fiber action potential model. A motor unit action potential was assumed to be the sum of the individual muscle fiber action potentials. A computational procedure, based on the notion of isopotential layers, was developed which substantially reduced the calculation time required to estimate motor unit action potentials. The simulations indicated that: 1) only those motor units with muscle fibers located within 10–12 mm of the electrodes would contribute significant signal energy to the surface EMG, 2) variation in surface area of electrodes has little effect on the detection depth of motor unit action potentials, 3) increased interelectrode spacing moderately increases detection depth, and 4) the frequency content of action potentials decreases steeply with increased electrode-motor unit territory distance.     

Development of homogeneous fast and slow motor units in the neonatal mouse soleus muscle

We studied the fiber type composition and contractile properties of mouse soleus motor units at 2 days, 5 days and 2 weeks of age. We used Lucifer Yellow injection to mark muscle fibers belonging to the same motor unit in the two youngest age groups, and the traditional method of glycogen depletion in the oldest. The age groups were chosen because 2 days is at the end of muscle fiber production; 5 days is at the start of synapse elimination in the muscle and 2 weeks is at the end. Muscle fibers were classified as fast (F) or slow (S) on the basis of their myosin heavy chain (MHC) content, as determined by different monoclonal antibodies. Motor units are already dominated by either F- or S-fibers at 2 days, suggesting an early preferential innervation of the two types of fibers. A substantial part of the remaining refinement of the innervation takes place during the next 3 days, while the total number of terminals in the muscle remains constant. This is most easily explained by an exchange of aberrant for correct synapses during this period. A smaller part of the refinement of the innervation occurs during the subsequent period of synapse elimination.     

Architecture of the hind limb muscles of cats: Functional significance

Force, velocity, and displacement properties of a muscle are determined in large part by its architectural design. The relative effect of muscle architecture on these physiological variables was studied by determining muscle weight, fiber length, average sarcomere length, and approximate angle of pinnation for 24 cat hind limb muscles. Muscle lengths ranged from 28.3 to 144 mm, whereas fiber lengths ranged from 8.4 to 105.5 mm. Generally, fiber to muscle length ratios were similar throughout a muscle. Estimated angles of pinnation of muscle fibers varied from 0 to 21° with most having an angle of less than 10°. The cross-sectional area of the knee extensors was similar to the knee flexors (16.43 vs. 16.83 cm²) whereas the cross-sectional area of the ankle extensors was more than six times greater than the ankle flexors (18.59 vs. 2.83 cm²). There was a 6.7-fold difference in the maximal force between muscles, when normalized to a constant weight, that could be attributed to architectural features. Rations of wet weight to predicted maximal tetanic tension for each muscle and group were calculated to compare the relative priority of muscle force versus muscle length-velocity for a given mass of muscle. These ratios varied from 0.4 to 4.84. The ratios suggest that velocity and/or displacement is a priority for the hamstrings, whereas force is a priority for the quadriceps and lower leg muscles. As much as a 12.6-fold difference in maximal velocity between muscles can be attributed to differences in fiber lengths. This can be compared to approximately a 2.5-fold difference in maximal velocity reported to occur as a result of biochemical (intrinsic) differences.