Brenner B
Basic Res Cardiol. 1979 Mar-Apr;74(2):177-202. doi: 10.1007/BF01907820.
The existence of still functioning vesicles of the sarcoplasmic reticulum (SR) in glycerinated rabbit psoas muscle fibers and cat myocardium is demonstrated by the use of activation (force development and shortening) of the contractile system as an indicator for "sarcoplasmic", free Ca++ concentration and by electron microscopic studies. Furthermore, the conditions for Ca++ release and uptake by the SR vesicles were analyzed. (1) As indicated by electron microscopic studies, glycerinated skeletal and heart muscle preparations contain SR vesicles (diameter 0.1--0.2 mu). Detergent treatment as used by Julian (1971) causes neither morphological change in the contractile system nor in the SR vesicles. (2) The delay in tension development of glycerinated preparations after an increase in free Ca++ concentration of the bathing medium is demonstrated to be caused at least partly by Ca++ uptake in these SR vesicles. (3) The SR vesicles of glycerinated preparations are able to accumulate Ca++ just at concentrations which are subthreshold for tension development. The accumulated Ca++ can be detected by a transient force development resulting from abolishment of the Ca++ storage capacity of the SR vesicles caused by high concentrations of detergents. (4) Ca++-induced Ca++ release is also demonstrated in glycerinated preparations. After Ca++ loading this release mechanism occurs at Ca++ concentrations in the bathing medium starting at about 10(-7,48) Mol/l both in skeletal and heart muscle preparations. (5) Quick stretch of Ca++-loaded, relaxed preparations caused local contractions of the myofibrils in both single fibers and fiber bundles of glycerinated rabbit psoas muscle. Skeletal fiber bundles and cat myocardium show a transient tension development parallel to these local contractions. These effects are demonstrated to be due to a stretch-induced Ca++ release from the SR vesicles. (6) Length-controlled quick stretches or quick releases of partly activated preparations cause sinusoidal tension oscillations superimposed on the well-known, delayed active tension development. In contrast, fully activated preparations do not show superimposed oscillations in tension. Force-controlled quick stretches or quick releases induce equivalent oscillations in length which are more distinct than the tension oscillations, however. These oscillations are also demonstrated to be due to stretch or release-induced changes in the Ca++ fluxes between SR vesicles and the "sarcoplasm". (7) This stretch induced release of Ca++ from the SR could be one reason for the multiple-phase course of active and passive isometric tension development in vivo following quick changes in length, and could also partly explain the prestretch dependence of tension development which cannot be explained from the degree of overlapping of the contractile filaments alone...
通过将收缩系统的激活(力的产生和缩短)作为“肌浆”中游离钙离子浓度的指标,并结合电子显微镜研究,证实了甘油处理的兔腰大肌纤维和猫心肌中肌浆网(SR)的囊泡仍具有功能。此外,还分析了SR囊泡释放和摄取钙离子的条件。(1)电子显微镜研究表明,甘油处理的骨骼肌和心肌标本含有SR囊泡(直径0.1 - 0.2微米)。Julian(1971)所采用的去污剂处理既未引起收缩系统的形态变化,也未引起SR囊泡的形态变化。(2)已证明,在浴液中游离钙离子浓度升高后,甘油处理标本张力发展的延迟至少部分是由这些SR囊泡摄取钙离子所致。(3)甘油处理标本的SR囊泡能够在低于产生张力阈值的浓度下积累钙离子。通过高浓度去污剂破坏SR囊泡的钙离子储存能力所导致的瞬时力的产生,可以检测到积累的钙离子。(4)在甘油处理的标本中也证实了钙离子诱导的钙离子释放。在钙离子加载后,这种释放机制在骨骼肌和心肌标本的浴液中钙离子浓度约为10^(-7.48)摩尔/升时开始出现。(5)对钙离子加载的松弛标本进行快速拉伸,导致甘油处理的兔腰大肌单纤维和纤维束中的肌原纤维发生局部收缩。骨骼肌纤维束和猫心肌表现出与这些局部收缩平行的瞬时张力发展。已证明这些效应是由于拉伸诱导SR囊泡释放钙离子所致。(6)对部分激活的标本进行长度控制的快速拉伸或快速释放,会在众所周知的延迟主动张力发展之上叠加正弦张力振荡。相比之下,完全激活的标本在张力上不显示叠加振荡。力控制的快速拉伸或快速释放会在长度上诱导等效振荡,然而,这些振荡比张力振荡更明显。这些振荡也被证明是由于拉伸或释放诱导的SR囊泡与“肌浆”之间钙离子通量的变化所致。(7)这种拉伸诱导的SR钙离子释放可能是体内长度快速变化后主动和被动等长张力发展呈现多相过程的一个原因,也可以部分解释仅从收缩细丝重叠程度无法解释的张力发展对预拉伸的依赖性……