Department of Zoology, La Trobe University, Melbourne, Victoria, Australia.
Biophys J. 2011 Jun 8;100(11):2688-97. doi: 10.1016/j.bpj.2011.04.027.
The stability of the filament lattice in relaxed striated muscle can be viewed as a balance of electrostatic and van der Waals forces. The simplest electrostatic model, where actin and myosin filaments are treated as charged cylinders, generates reasonable lattice spacings for skinned fibers. However, this model predicts excessive radial stiffness under osmotic pressure and cannot account for the initial pressure (∼1 kPa) required for significant compression. Good agreement with frog compression data is obtained with an extended model, in which S1 heads are weakly attached to actin when the lattice spacing is reduced below a critical value; further compression moves fixed negative charges on the heads closer to the myofilament backbone as they attach at a more acute angle to actin. The model predicts pH data in which the lattice shrinks as pH is lowered and protons bind to filaments. Electrostatic screening implies that the lattice shrinks with increasing ionic strength, but the observed expansion of the frog lattice at ionic strengths above 0.1 M with KCl might be explained if Cl(-) binds to sites on the motor domain of S1. With myosin-myosin and actin-actin interactions, the predicted lattice spacing decreases slightly with sarcomere length, with a more rapid decrease when actin-myosin filament overlap is very small.
在松弛的横纹肌中,丝状体晶格的稳定性可以看作是静电力和范德华力的平衡。最简单的静电模型将肌动蛋白和肌球蛋白丝处理为带电圆柱,为去皮纤维生成合理的晶格间距。然而,该模型预测在渗透压下会产生过大的径向刚度,并且无法解释用于显著压缩所需的初始压力(约 1kPa)。通过扩展模型可以很好地与青蛙压缩数据相吻合,在该模型中,当晶格间距减小到低于临界值时,S1 头部与肌动蛋白的弱结合;进一步的压缩使头部上的固定负电荷更靠近肌球蛋白丝主干,因为它们以更锐角附着在肌动蛋白上。该模型预测了晶格随 pH 值降低和质子结合到丝状体而收缩的 pH 值数据。静电屏蔽意味着晶格随离子强度的增加而收缩,但如果 Cl(-)结合到 S1 的马达结构域上,则可以解释在离子强度高于 0.1M 的 KCl 中观察到的青蛙晶格的扩张。对于肌球蛋白-肌球蛋白和肌动蛋白-肌动蛋白相互作用,预测的晶格间距随肌节长度略有减小,当肌动蛋白-肌球蛋白丝重叠非常小时,减小速度更快。