Nonuniform changes in fibre types in the soleus muscle of the developing rat

Muscle fibre composition among the proximal (25%), middle (50%) and distal (75%) regions of the rat soleus muscle at various ages were compared to investigate whether the region-specific changes in fibre types known to occur under nonphysiological conditions (i.e. electrical stimulation along with immobilization in a lengthened position) also occur in the developing muscle. In addition, we attempted to detect fibres with nonuniform histochemical properties in the different segments, processing with myofibrillar actomyosin adenosine triphosphatase after pre-incubation at pH 10.3 against successive cryosections (200 m apart). Samples were obtained from 66 Wistar rats of both sexes ranging in age from 13 to 85 days and subdivided into age groups of 2, 3, 4, 5 and 12 weeks. The mass and length of the soleus muscle increased most significantly at age 3–4 and 2–4 weeks, respectively. The distal region had a significantly lower percentage of type I fibres than the middle region at age 3 weeks, than the proximal and middle regions at age 4 weeks. In addition, some fibres [0.7 (SD 0.5)% n=19 ± 13] possessing nonuniform histochemical properties in different segments were observed from the middle and distal region at age 4 weeks. These findings would suggest that transformations from type II to type I fibres in the soleus muscle of the developing rat occur from the proximal or middle segments.     

Venous drainage of the soleus muscle

Soleus veins have been implicated as the site for deep venous thrombosis initiation. Detailed anatomic knowledge is required for the early diagnosis using non-invasive ultrasound techniques. In the present work, we describe the anatomy of the veins that emerge from the ventral face of the soleus muscle. Twenty-eight soleus muscles were dissected and 543 veins were found. The number of veins per leg ranged from 7 to 38. The distribution of these veins per quadrant ranged from 0 to 12. The greatest number of veins occurred at the superior lateral quadrant. Most soleus veins mainly drained into the posterior tibial and fibular veins. The mean length of the soleus veins ranged from 0.907 to 2.804 cm. We conclude that there is a wide variability in the distribution of soleus veins through the soleus muscle and their quadrants. The majority of the soleus veins drain into the tibial and fibular veins.     

The time course of thyroid-hormone-induced changes in the isotonic and isometric properties of rat soleus muscle

Rat thyroidectomy resulted in changes in a number of parameters used to characterise the mechanical and histochemical status of skeletal muscle. Thus thyroidectomy resulted in a prolongation of soleus slow-twitch muscle isometric contraction time and half-relaxation time with a reduced maximum velocity of shortening and maximum rate of development of tetanic tension but no significant change in twitch: tetanus ratio i.e. the ratio of twitch force/unit area to tetanic force/unit area. In addition the percentage of IIA fibres was reduced and the percentage of type I fibres increased. Triiodothyronine, administered to hypothyroid rats, brought about a speeding of these parameters again with no change in twitch: tetanus ratio. There was an increase in the percentage of IIA fibres with a concomitant reduction in the percentage of type I fibres. These changes were induced over 18 days and resulted in isotonic and isometric properties almost identical to those of soleus muscles from chronically hyperthyroid rats; speeding could be detected as early as 2 days after triiodothyronine had been given. Consideration is given to the possibility that changes in myosin isoforms and/or the kinetics of changes in intracellular calcium concentration in activation and relaxation could account for the time course of the observed changes.     

Activity‐induced fiber regeneration in rat soleus muscle

In an attempt to understand why muscle recovery is limited following atrophy due to limb immobilization, satellite cell activity and muscle fiber regeneration were analyzed in rat soleus muscles. Adult rat hindlimbs were immobilized in plaster casts for a period of two to ten weeks. Soleus muscles were examined by electron microscopy for evidence of fiber degeneration or regeneration, and to quantify satellite cell nuclei. Immunocytochemical localization of embryonic myosin was used to identify regenerating myofibers. Soleus muscle wet weight to body weight ratios for the casted muscles significantly decreased over the 10‐week immobilization period. The casted muscles displayed ultrastructural evidence of minor fiber damage, including myofibrillar atrophy, Z‐disc disruption, and abnormal triadic junctions. No ultrastructural evidence of regeneration was seen in the casted animals. The number of satellite cells in the casted muscles significantly decreased from 6.4% to 3.3% by eight to 10 weeks of immobilization. Approximately 1.0% of extrafusal fibers in the control soleus muscles appeared to be regenerating since they expressed embryonic myosin and were of a small diameter, while in casted muscles, only 0.1% of the fibers were embryonic myosin‐positive. Following release from immobilization, a reappearance of embryonic myosin‐positive fibers was noted within four days of renewed activity. In contrast to control muscles, embryonic myosin‐positive fibers in the recovery muscles included both small and large diameter fibers. Subtle changes in functional activity influence muscle damage and subsequent myofiber regeneration. Reduced activity reduces muscle fiber regeneration, while increased activity, as seen by increased hindlimb weight bearing and return to normal activity following immobilization, increase regenerating fibers and also the expression of embryonic myosin in adult fibers. Anat Rec 258:176–185, 2000. © 2000 Wiley‐Liss, Inc.     

Some observations on the efferent innervation of rat soleus muscle spindles

Summary In the small segmental tail muscles of the rat beta fibres provide exclusively the dynamic fusimotor control, while gamma fibres provide exclusively the static fusimor control. The present experiments were made to investigate the fusimotor innervation of spindles in a large muscle of the rat, the soleus, and thus to determine the occurrence and significance of beta innervation in this muscle. Our results have revealed no case of beta innervation in the rat soleus. As a consequence of our experimental method, however, we would not claim that beta innervation does not exist in the soleus, only that it must play an insignificant role relative to that seen in the tail segmental muscles. Investigations of the fusimotor innervation of eight spindles were sufficiently complete to warrant detailed illustration. The number of gamma fibres ranged from two to four. In every case the slowest conducting gamma fibre was dynamic. However, the conduction velocity spectra for the static and dynamic gamma fibres to rat soleus overlap to such an extent that it is impossible to use conduction velocity as the sole guide to functional gamma fibre classification. The pooled results from the eight spindles fully investigated provide a ratio of static to dynamic gamma fibres of approximately 1:1. Other evidence discussed in the paper suggests that in the muscle nerve the ratio is considerably higher. These differences are reconciled if the dynamic gamma fibres branch more profusely and innervate more spindles than do the static gamma fibres.     

Effect of ischemia on contractile and histochemical properties of the rat soleus muscle

Ligature and section of the abdominal aorta results in only minor and temporary functional and metabolic changes in the slow soleus muscle of the rat. A very small decrease in maximal tetanic tension corresponds to a few scattered areas of damaged and necrotic muscle fibres, in which decreased succinic dehydrogenase and loss of phosphorylase activity was observed.A new experimental approach, i.e. ligature and section of the abdominal aorta combined with terminal devascularisation, preserving intact tendons and innervation of the muscle causes maximal muscle ischemia, followed by an almost complete loss of tetanic tension output, marked shortening of contraction time and profound morphological and histochemical changes. The decrease in succinic dehydrogenase and ATPase activities and loss of phosphorylase activity occur in the majority of degenerating muscle fibres except for a thin rim of peripheral fibres during the first 4 days. Subsequently, the contractile properties recover gradually and enzyme activities reappear in the regenerating muscle fibres simultaneously with new revascularisation. Thirty days after the operation all the parameters observed returned to control values.Deceased August 6, 1977     

Effect of a single session of electrical stimulation on activity and expression of citrate synthase and antioxidant enzymes in rat soleus muscle

The aim of our study was to investigate the effect of a single high intensity session of muscle contractions on the activity and expression of citrate synthase (CS) and of the following major antioxidant enzymes: Mn-superoxide dismutase (Mn-SOD), Cu,Zn-superoxide dismutase (Cu,Zn-SOD), catalase (CAT), and glutathione peroxidase (GPX). To accomplish this, soleus muscles of male Wistar rats were subjected to contractions using a intense electrical stimulation (ES) protocol. Soleus muscles were isolated either immediately or 1 h after the contractions and utilized for enzyme activity determination, and for analysis of gene expression by quantitative PCR. A significant increase in maximal activity (63%) and expression (80%) of CS was observed in stimulated soleus muscles, isolated 1 h after ES as compared to controls. However, this effect was not observed in muscles isolated immediately after ES. By using macroarray and Real Time RT-PCR analysis, an increase in expression of Mn-SOD, Cu,Zn-SOD, CAT, and GPX was also found. Interestingly, of these enzymes, only CAT activity was significantly increased (44%) 1 h after ES in soleus muscle. These results indicate that acute ES up-regulates activity and expression of CS and CAT in soleus muscles. This increase in expression of CAT may play an important role in counteracting the potential deleterious effects of elevated oxidative stress induced by a high oxidative demand in skeletal muscles subjected to exercise training.     

Stiffness changes and fibre type transitions in rat soleus muscle produced by jumping training

Rat soleus muscles were overloaded with intent to induce a relative increase in fast fibres and modifications in muscular stiffness. The overloading technique was a training period consisting of an 11-week vertical jump programme. The method of controlled releases was used to obtain tension/extension curves characterizing the elastic behaviour of the soleus. Fibre typing was made by myofibrillar adenosine 5-triphosphatase staining. With regard to a control group, training resulted in a relative decrease in type I fibres for the benefit of type II fibres. Training also induced a decrease in muscle stiffness as attested notably by significant differences in maximal extension. These results are interpreted in terms of modifications occurring in the active fraction of the so-called series elastic component.     

缺氧及缺氧-复合运动大鼠比目鱼肌肌球蛋白重链组成变化

 目的： 观察缺氧及缺氧复合运动条件下大鼠比目鱼肌肌球蛋白重链(MHC)异构体组成的变化。方法: Wistar大鼠随机分为4组：平原对照组、缺氧组、平原运动组和缺氧复合运动组。缺氧复合运动组大鼠持续暴露于模拟海拔5 000 m高原5周，每天降至4 000 m高原进行游泳运动1 h（6 d/week），运动结束后回升至5 000 m；缺氧组大鼠同时在低压舱内相同海拔高度饲养，但不进行游泳运动；平原运动组和平原对照组在舱外同时饲养，其中平原运动组每天进行游泳运动1 h（6 d/week）。在末次运动结束后24 h处死大鼠，分离后肢比目鱼肌。用含30%甘油的SDS-PAGE电泳分离比目鱼肌MHC异构体，观察MHC异构体组成变化。结果: 平原游泳运动大鼠比目鱼肌重量指数增加，而缺氧复合运动组与平原对照比较无显著差异；大鼠比目鱼肌中段MHC主要由Ⅰ(78％)、Ⅱa(22％)2种异构体组成，缺氧组比目鱼肌MHC由Ⅱa向Ⅰ型转变，单纯运动和缺氧复合运动组与缺氧组变化趋势相同，但缺氧复合运动组显著低于单纯运动组。结论: 慢性缺氧和运动训练后，大鼠比目鱼肌肌纤维MHC组成向能量利用效率更高的慢收缩肌纤维Ⅰ型转变，提示慢性缺氧和缺氧复合运动后比目鱼肌能量利用效率增加。     

Compensatory effects of chronic electrostimulation on unweighted rat soleus muscle

The purpose of this study was to investigate the effects of electrostimulation in counteracting the transformation of the unweighted rat soleus muscle. The stimulation resembled the firing patterns of normal slow motor units and was imposed during hindlimb suspension. For the 10-day hindlimb suspended rats, the transformation of the slow soleus muscle towards a faster type was characterized by a decrease in the time to peak tension and the half-relaxation time of the twitch, a reduction in the P ₂₀/P ₀ index, i. e. the ratio of the subtetanic tension at 20 Hz relative to the tetanic tension, and a decrease in the percentage distributions of type I fibres accompanied by an increase of type IIa and IIc fibres. These changes were prevented by electrostimulation since, for the parameters mentioned above, no significant difference was observed in the soleus of the suspended rats that received electrostimulation when compared with the control rats. Nevertheless, neither the loss of mass nor the decrease in force output in the suspended rats were prevented by electrostimulation. The present results suggest a positive compensation of the suspension-induced alterations in the contractile and histochemical properties of the soleus muscle by means of chronic electrostimulation, which, however, do not prevent atrophy or the loss of contractile force.     

Relationship between myosin heavy chain IId isoform and fibre types in soleus muscle of the rat after hindlimb suspension

Summary The relationship between the myosin heavy chain (HC) IId isoform and histochemically defined fibre types was investigated in the rat soleus muscle after hindlimb suspension. After 4 weeks of suspension, right and left muscles were removed and fibre type composition and total fibre number were examined by histochemical myosin adenosine triphosphatase staining sections. Myosin HC isoforms were analysed by sodium dodecyl sulphate polyacrylamide gel electrophoresis. After the suspension, there was a significant decrease in the percentage of type I fibres and a concomitant increase in that of type IIa fibres. However, the total number of fibres was not affected by suspension. The synthesis of HC IId isoform, which was not found in the control, and the decrease in the ratio of slow type myosin heavy chain isoform (HC I) were observed after suspension. These results would may suggest that the change of fibre type composition was caused by a shift from type I to IIa fibres after suspension. Furthermore, it could be suggested that the synthesis of HC IId isoform occurred during the stage of type shift from type I to IIa fibres.     

Effects of hypobaric hypoxia on histochemical fibre-type composition and myosin heavy chain isoform component in the rat soleus muscle

Histochemical fibre-type composition and myosin heavy chain isoform component in the soleus muscle were studied in normoxic rats at postnatal ages of 5, 10, 15, and 20 weeks and in rats exposed to hypobaric hypoxia (460 torr) for 5 weeks from postnatal ages of 5, 10, and 15 weeks. The increase in the percentage of type I fibres and the concomitant decrease in that of type IIa fibres in the soleus muscle of normoxic rats were observed until 15 weeks of age. On the other hand, no change in the fibre-type composition of the muscle during postnatal development was observed in hypoxic rats, irrespective of the age at which they were exposed to hypoxia. The changes in the myosin heavy chain isoform component (MHC I and MHC IIa) of the muscle during postnatal development and by hypoxia corresponded well with those in the muscle fibre-type composition. It is concluded that hypobaric hypoxia inhibits the growth-related shift of muscle fibre-types from type IIa to type I and of myosin heavy chain isoforms from MHC IIa to MHC I in the rat soleus muscle, and that there are no changes in the muscle fibre-type composition or the myosin heavy chain isoform component caused by hypoxia after the shifts in these parameters which occur during postnatal development are completed.     

Enhanced stretch reflex excitability of the soleus muscle in experienced swimmers

The purpose of this study was to investigate effects of long-term participation to swimming on adaptations of spinal reflex excitability. To this end, mechanically induced stretch reflex (SR) and electrically induced Hoffmann (H-) reflex of the soleus muscle were investigated between swimmers with experience of more than 10 years and non-trained individuals while sitting at rest. The amplitude and the gain (stretch velocity vs. amplitude of the reflex response) of the SR were significantly greater in the swimming group than in the non-trained control group. Similarly, the responses of the H-reflex were also significantly greater in the swimming group than in the non-trained control group. Results of this study demonstrated that the spinal reflex excitability in experienced swimmers was far more enhanced than in non-trained individuals.     

Structure-function relationship of soleus muscle fibres from the rhesus monkey

Functional and structural properties of rhesus monkey skinned fibres were studied in order to examine the relationship between calcium/strontium (Ca/Sr) activation characteristics and protein composition. The fibres were classified according to their Ca/Sr affinity into slow (61%) and fast groups (39%). According to the myosin isoform composition, two additional hybrid types were defined. Thus, four profiles were characterized: two corresponding to slow (S) and fast (F) isoforms and two corresponding to a mixed proportion of slow and fast isoforms. They were called hybrid slow (HS) or hybrid fast (HF) based on the predominant myosin isoform. Tension/pCa parameters and maximal shortening velocities were determined. S fibres showed a higher pCa threshold and affinity as well as shallower slopes of their tension/pCa curve than did F fibres. HS and HF fibres exhibited tension/pCa curves which were positioned close to those of S and F fibres, respectively. No significant difference was observed between S and HS fibres or between F and HF fibres. Maximal shortening velocity values were higher for fibres expressing predominantly fast myosin isoforms. We suggest than when both S and F isoforms of myofibrillar proteins are expressed in a muscle fibre, the functional properties are mainly governed by the predominant isoform.     

腓血管蒂比目鱼肌皮瓣逆行转位修复术的应用解剖

目的：为以腓血管为蒂比目鱼肌皮瓣逆行转位修复小腿中、下段组织缺损提供解剖学基础。方法：在３３侧经动脉灌注乳胶的下肢标本及６侧动脉铸型下肢标本上,解剖观测比目鱼肌形态,腓动脉比目鱼肌支的分布及腓动脉与胫前、后动脉之间的吻合。结果：腓动脉发出１～４条比目鱼肌支,外径２．０±０．５（１．０～３．８）ｍｍ,由其发出至皮肤的血管在比目鱼肌内或沿肌间隔穿行,穿出点在腓骨头下９．９±３．０ｃｍ,分布范围１６ｃｍ×９ｃｍ。腓动脉下端与胫前、后动脉间有丰富的吻合支和粗大的交通支,最近端交通支距内、外踝连线５．５±０．８（３．６～７．４）ｃｍ,外径１．２±０．３ｍｍ。结论：以腓血管为蒂可以形成比目鱼肌皮瓣;该瓣逆行转位可修复小腿中下段缺损,具有血供可靠、损伤小、操作简单的优点     

Influence of chronic stretching upon rat soleus muscle during non-weight-bearing conditions

Morphological, contractile and histochemical properties as well as the myosin heavy chain (MHC) composition of rat soleus muscles were studied after 14 days of non-weight-bearing (NWB) and after immobilization of the foot in dorsiflexion of NWB rats. Significant reductions in soleus mass, fibre sizes and tetanic tension were found after 14 days of NWB. Furthermore, a transformation of the slow-twitch soleus muscle towards a faster type was characterized by a decrease in twitch time parameters, an increase in the fast-twitch type IIA fibre proportion and an increase in fast-twitch type MHC isoforms. Our results showed that the immobilization of the soleus muscle in a lengthened position during NWB not only prevented the loss of muscular mass and force output, but also counteracted the slow to faster shift in contractile and phenotypical parameters normally associated with NWB conditions.     

Changes in fibre-type composition and myosin heavy-chain IId isoform in rat soleus muscle during recovery period after hindlimb suspension

We examined the changes in myosin heavy-chain (HC) isoforms and fibre-type composition in rat soleus muscle using both myosin adenosine triphosphatase staining and sodium dodecyl sulphate/polyacrylamide gel electrophoresis (SDS-PAGE) analyses during the recovery period after 4 weeks of hindlimb suspension. Although there was no change in type IIc fibres after the suspension, an increase in this type of fibres was observed during the 1- to 4-week recovery period. The increase in type Ilc fibres was considered to be due to a shift from type Ila to IIc fibres. The SDS-PAGE analysis revealed the presence of the HC IId isoform, which was not observed in the control muscle, after a 4-week hindlimb suspension. The HC IId isoform gradually decreased over 3 weeks of recovery and disappeared in the 4th week of recovery after the suspension. These results suggest that the hypogravity conditions induced by hindlimb suspension stimulated the synthesis of the HC IId isoform, whereas an increase in mechanical load to the muscle accelerated the degradation of the HC IId isoform and the synthesis of type Ilc fibres during the recovery period after hindlimb suspension.     

压力和尾吊对大鼠骨骼肌生长的影响

目的建立尾吊和压力大鼠实验模型,比较压力与尾吊对大鼠骨骼肌生长的不同影响。方法 36只雄性SD大鼠随机分为3组:正常对照组、尾吊组和压力组,每组分两个阶段(7、14 d)进行观察。实验结束后测量比目鱼肌与趾长伸肌的湿重体重比、肌纤维横截面积和直径,以及血清中IGF-1浓度。结果压力作用7 d与尾吊7 d后,比目鱼肌湿重体重比、肌纤维横截面积、肌纤维直径都较对照组显著减少(P<0.05);压力组分别减少23.52%、14.26%、13.47%(P<0.05),尾吊组分别减少23.52%、33.07%、25.09%(P<0.05)。压力作用14 d后,各指标分别减少20.51%、-10.49%、-5.73%,都低于7 d压力组的减少量(P<0.05);而尾吊14 d后,比目鱼肌湿重体重比减少了46.15%,显著高于7 d尾吊组的减少量。血清IGF-1浓度和趾长伸肌的改变在压力、尾吊组间没有显著性差异。结论压力对比目鱼肌生长影响的过程不同于尾吊。压力作用初期是以炎症反应为主的组织损伤过程,待肌细胞适应压力环境后,可能产生一定的功能适应性生长。因此,在临床上无论假肢接受腔设计还是康复训练,考虑接受腔压力对内部肌肉损伤的影响,将有助于对残端肌肉组织的保护。     

Effects of immobilization on the rat soleus muscle in relation to age

A hind limb of young adult, adult and old male Wistar rats (4–5, 6–7 and 20–21 months, respectively) was immobilized for 4 weeks by a plaster cast with the knee and ankle joints in a resting position. Enzyme-histochemical, morphometrical and contractile characteristics of the soleus muscle were compared with those in age-matched controls. A pronounced decrease in muscle mass and cross-sectional muscle fibre area was found at all ages. The degree of atrophy after immobilization did not differ between different fibre types in each age group, but the decrease in fibre area was less pronounced in old animals (i.e. the fibre area was decreased by 49–64, 53–66 and 27–38% in young adult, adult and old animals, respectively). The maximum tetanus force was decreased in all age groups (by 73, 78 and 69% in young adult, adult and old rats, respectively) as was the tetanus tension (i.e. tetanus force divided by muscle fibre cross-sectional area). The contraction time of the isometric twitch was significantly altered, i.e. decreased, only in the youngest age group, although it also tended to decrease in old age. A significant increase in the number and proportion of fibre types intermediate to types I and 11 A, was found in the immobilized muscle of 4–5- and 6–7-month-old animals, but not in that of old ones (i.e. the proportion of intermediate fibres increased by 14, 13 and 2% in young adult, adult and old animals, respectively). Thus, in contrast to the atrophic changes, the contractile alterations after immobilization were not markedly different between young and old age. It is further concluded that the age-related fast-to-slow muscle fibre transition that occurs in normal soleus during maturation and growth can be partly reversed by restrictions of the normal muscle activity and that the ability of the soleus to modulate its fibre-type composition in response to a change in activity may be diminished in old age.     

Different effects of physical training on the morphology of motor nerve terminals in the rat extensor digitorum longus and soleus muscles

Summary The morphology of nerve terminals in the rat extensor digitorum longus and soleus muscles was studied with light microscopy in 13-week-old male animals after 6 weeks of treadmill running and compared with data from untrained controls. The terminals were stained with methylene blue. Physical training tended to increase the area and length of the nerve terminals in relation to the corresponding muscle fiber diameter, and to reduce the density of nerve terminal varicosities, but significant differences between the trained group and the control group were obtained only in the extensor digitorum longus muscle. The different degrees of effect on the nerve terminals in the two muscles may be due to different abilities to respond to the training, but may also be due to differences in work load caused by the training. The effect of training on extensor digitorum longus junctions may reflect some transformation from fast to slow morphological characteristics.