Callister Robert J, Pierce Patricia A, McDonagh Jennifer C, Stuart Douglas G
School of Biomedical Sciences, Faculty of Health, University of Newcastle, Callaghan, NSW 2308, Australia.
J Morphol. 2005 Apr;264(1):62-74. doi: 10.1002/jmor.10318.
A description is provided of the ratio of slow-tonic vs. slow- and fast-twitch fibers for five muscles in the adult turtle, Pseudemys (Trachemys) scripta elegans. The cross-sectional area of each fiber type and an estimation of the relative (weighted) cross-sectional area occupied by the different fiber types are also provided. Two hindlimb muscles (flexor digitorum longus, FDL; external gastrocnemius, EG) were selected on the basis of their suitability for future motor-unit studies. Three neck muscles (the fourth head of testo-cervicis, TeC4; the fourth head of retrahens capitus collique, RCCQ4; transversalis cervicis, TrC) were chosen for their progressively decreasing oxidative capacity. Serial sections were stained for myosin adenosine triphosphatase (ATPase), NADH-diaphorase, and alpha-glycerophosphate dehydrogenase (alpha-GPDH). Conventional fiber-type classification was then performed using indirect markers for contraction speed and oxidative (aerobic) vs. glycolytic (anaerobic) metabolism: i.e., slow oxidative (SO, including slow-twitch and possibly slow-tonic fibers), fast-twitch, oxidative-glycolytic (FOG), and fast-twitch glycolytic (Fg) fibers. Slow-tonic fibers in the SO class were then revealed by directing the monoclonal antibody, ALD-58 (raised against the slow-tonic fiber myosin heavy chain of chicken anterior latissimus dorsi), to additional muscle cross sections. All five of the tested muscles contained the four fiber types, with the ATPase-stained fibers including both slow-tonic and slow-twitch fibers. The extreme distributions of SO fibers were in the predominately glycolytic TrC vs. the predominately oxidative TeC4 muscle (TrC-SO, 9%; FOG, 20%; Fg, 71% vs. TeC4-SO, 58%: FOG, 16%; Fg, 25%). Across the five muscles, the relative prevalence of slow-tonic fibers (4-47%) paralleled that of the SO fibers (9-58%). TeC4 had the highest prevalence of slow-tonic fibers (47%). The test muscles exhibited varying degrees of regional concentration of each fiber type, with the distribution of slow-tonic fibers paralleling that of the SO fibers. In the five test muscles, fiber cross-sectional area was usually ranked Fg > FOG > SO, and slow-twitch always > slow-tonic. In terms of weighted cross-sectional area, which provides a coarse-grain measure of each fiber type's potential contribution to whole muscle force, all five muscles exhibited a higher Fg and lower SO contribution to cross-sectional area than suggested by their corresponding fiber-type prevalence. This was also the case for the slow-twitch vs. slow-tonic fibers. We conclude that slow-tonic fibers are widespread in turtle muscle. The weighted cross-sectional area evidence suggested, however, that their contribution to force generation is minor except in highly oxidative muscles, with a special functional role, like TeC4. There is discussion of: 1) the relationship between the present results and previous work on homologous neck and hindlimb muscles in other nonmammalian species, and 2) the potential motoneuronal innervation of slow-tonic fibers in turtle hindlimb muscles.
本文描述了成年秀丽锦龟(Pseudemys (Trachemys) scripta elegans)五种肌肉中慢张力纤维与慢肌纤维和快肌纤维的比例。还提供了每种纤维类型的横截面积以及不同纤维类型所占相对(加权)横截面积的估计值。根据其对未来运动单位研究的适用性,选择了两块后肢肌肉(趾长屈肌,FDL;腓肠外侧肌,EG)。选择了三块颈部肌肉(颈睾提肌第四头肌,TeC4;颈斜头肌第四头肌,RCCQ4;颈横肌,TrC),因为它们的氧化能力逐渐降低。连续切片用肌球蛋白三磷酸腺苷酶(ATPase)、NADH-黄递酶和α-甘油磷酸脱氢酶(α-GPDH)染色。然后使用收缩速度和氧化(有氧)与糖酵解(无氧)代谢的间接标记物进行传统的纤维类型分类:即慢氧化(SO,包括慢肌纤维和可能的慢张力纤维)、快肌纤维、氧化糖酵解(FOG)和快肌糖酵解(Fg)纤维。然后通过将单克隆抗体ALD-58(针对鸡背阔肌前肌的慢张力纤维肌球蛋白重链产生)应用于额外的肌肉横截面,揭示SO类中的慢张力纤维。所有五块测试肌肉都包含这四种纤维类型,ATPase染色的纤维包括慢张力纤维和慢肌纤维。SO纤维的极端分布情况为:在主要进行糖酵解的TrC肌肉中,SO纤维占9%,FOG纤维占20%,Fg纤维占71%;而在主要进行氧化代谢的TeC4肌肉中,SO纤维占58%,FOG纤维占16%,Fg纤维占25%。在这五块肌肉中,慢张力纤维的相对比例(4 - 47%)与SO纤维的比例(9 - 58%)平行。TeC4中慢张力纤维的比例最高(47%)。测试肌肉中每种纤维类型呈现出不同程度的区域集中分布,慢张力纤维的分布与SO纤维平行。在这五块测试肌肉中,纤维横截面积通常排序为Fg > FOG > SO,且慢肌纤维总是大于慢张力纤维。就加权横截面积而言,它提供了每种纤维类型对整块肌肉力量潜在贡献的粗略衡量指标,所有五块肌肉中,Fg对横截面积的贡献高于其相应纤维类型比例所表明的,而SO的贡献则低于相应比例,慢肌纤维与慢张力纤维的情况也是如此。我们得出结论,慢张力纤维在龟类肌肉中广泛存在。然而,加权横截面积证据表明,除了像TeC4这样具有特殊功能作用的高氧化肌肉外,它们对力量产生的贡献较小。文中还讨论了:1)本研究结果与之前关于其他非哺乳动物物种同源颈部和后肢肌肉研究工作的关系,以及2)龟类后肢肌肉中慢张力纤维潜在的运动神经元支配情况。