Department of Physics, Faculty of Science, Tohoku University, Aoba, Sendai, Japan.
Prog Biophys Mol Biol. 2011 May;105(3):199-207. doi: 10.1016/j.pbiomolbio.2010.12.003. Epub 2010 Dec 13.
It is widely accepted that muscle cells take either force-generating or relaxing state in an all-or-none fashion through the so-called excitation-contraction coupling. On the other hand, the membrane-less contractile apparatus takes the third state, i.e., the auto-oscillation (SPOC) state, at the activation level that is intermediate between full activation and relaxation. Here, to explain the dynamics of all three states of muscle, we construct a novel theoretical model based on the balance of forces not only parallel but also perpendicular to the long axis of myofibrils, taking into account the experimental fact that the spacing of myofilament lattice changes with sarcomere length and upon contraction. This theory presents a phase diagram composed of several states of the contractile apparatus and explains the dynamic behavior of SPOC, e.g., periodical changes in sarcomere length with the saw-tooth waveform. The appropriate selection of the constant of the molecular friction due to the cross-bridge formation can explain the difference in the SPOC periods observed under various activating conditions and in different muscle types, i.e., skeletal and cardiac. The theory also predicts the existence of a weak oscillation state at the boundary between SPOC and relaxation regions in the phase diagram. Thus, the present theory comprehensively explains the characteristics of auto-oscillation and contraction in the contractile system of striated muscle.
人们普遍认为,肌肉细胞通过所谓的兴奋-收缩耦联以全或无的方式呈现出产生力或放松的状态。另一方面,无膜收缩装置在激活水平上处于第三种状态,即自动振荡(SPOC)状态,该激活水平介于完全激活和放松之间。在这里,为了解释肌肉的所有三种状态的动力学,我们构建了一个新的理论模型,该模型不仅基于平行于肌原纤维长轴的力的平衡,还基于垂直于肌原纤维长轴的力的平衡,同时考虑了实验事实,即肌节长度和收缩时肌丝晶格的间距会发生变化。该理论呈现出由收缩装置的几个状态组成的相图,并解释了 SPOC 的动态行为,例如,肌节长度的周期性变化呈锯齿波形状。由于横桥形成而导致的分子摩擦力常数的适当选择,可以解释在各种激活条件下观察到的 SPOC 周期以及不同肌肉类型(骨骼和心脏)之间的 SPOC 周期的差异。该理论还预测了在相图中 SPOC 和放松区域之间的边界处存在弱振荡状态。因此,本理论全面解释了有条纹肌肉收缩系统中自动振荡和收缩的特征。