Trombitás K, Baatsen P, Schreuder J, Pollack G H
Center for Bioengineering, University of Washington, Seattle 98195.
J Muscle Res Cell Motil. 1993 Dec;14(6):573-84. doi: 10.1007/BF00141554.
Although X-ray diffraction measurements imply almost constant filament separation during isometric contraction, such constancy does not hold at the level of the isolated cell; cell cross-section increases substantially during isometric contraction. This expansion could arise from accumulation of water drawn from other fibre regions, or from water drawn into the cell from outside. To distinguish between these hypotheses, we froze single fibres of frog skeletal muscle that were jacketed by a thin layer of water. Frozen fibres were freeze-substituted, sectioned transversely, and examined in the electron microscope. In fibres frozen during contraction, we found large amounts of water just beneath the sarcolemma, less in deeper regions, and almost none in the fibre core. Such gradients were absent or diminished in fibres frozen in the relaxed state. The water was not confined to the myofibril space alone; we found large water spaces between myofibrils, particularly near mitochondria. Accumulation of water between myofibrils and around mitochondria implies that the driving force for water movement probably lies outside the filament lattice, and may therefore be osmotic. The fact that the distribution was nonuniform-highest near the sarcolemma and lowest in the core--implies that the water was likely drawn from the thin jacket surrounding the cell. Thus, the contractile cycle appears to be associated with water entry into and exit from the cell.
尽管X射线衍射测量表明,在等长收缩过程中细丝间距几乎保持恒定,但在分离细胞水平上这种恒定并不成立;在等长收缩过程中细胞横截面积会大幅增加。这种扩张可能源于从其他纤维区域吸取的水的积累,或者源于从细胞外部吸入细胞的水。为了区分这些假设,我们冷冻了包裹着一层薄水层的青蛙骨骼肌单纤维。将冷冻的纤维进行冷冻置换,横向切片,并在电子显微镜下检查。在收缩过程中冷冻的纤维中,我们发现肌膜下方有大量的水,较深区域的水较少,而纤维核心几乎没有水。在松弛状态下冷冻的纤维中,这种梯度不存在或减小。水并不局限于肌原纤维空间;我们在肌原纤维之间发现了大的水空间,特别是靠近线粒体的地方。肌原纤维之间和线粒体周围水的积累意味着水移动的驱动力可能存在于细丝晶格之外,因此可能是渗透压。分布不均匀这一事实——在肌膜附近最高,在核心最低——意味着水可能是从围绕细胞的薄水层中吸取的。因此,收缩周期似乎与水进出细胞有关。