Tregear R T
AFRC Institute of Animal Physiology, Cambridge, UK.
Adv Exp Med Biol. 1988;226:513-26.
In view of the current disagreement about the degree of tension maintenance in rabbit fibres in AMPPNP, I have restudied the mechanics of glycerol-extracted rabbit psoas and Lethocerus flight muscle in rigor and in MgAMPPNP. Insect fibres elongated reversibly when AMPPNP was added; the effect required little nucleotide. Rabbit fibres showed only a relatively small reversible elongation on adding AMPPNP; my experimental evidence for mechanochemical equilibrium is therefore only certain for the insect muscle. At low muscle tension extension of either muscle type produced little tension decay; the stress-relaxation seen ('visco-elasticity') followed a power law of low order. At greater tension both muscles showed a second form of tension decay ('yield') which dominated at high tension, causing within a few minutes a near-total loss of added tension due to stretch. Below this tension the isometric tension decay was relatively slow, and considerable tension remained overnight. Yield dominated at a much lower tension in AMPPNP than in rigor, but relative to this changed level the speed of the yield process did not appear to alter greatly; again below a certain level tension decay became very slow. The effect of AMPPNP on yield was reversible and appeared similar in the two tissues. No obvious change occurred in any of these properties when the AMPPNP was purified before use or when a myokinase inhibitor and ADP-removing enzyme system were added. Thus in both preparations tension is maintained for a time long relative to the expected lifetime of an individual crossbridge, and its decay during yield is not accelerated by shortening that lifetime. These phenomena indicate that some process other than crossbridge detachment is limiting crossbridge slip; I am seeking an explanation in terms of interaction between neighbouring crossbridges.
鉴于目前对于在AMPPNP存在下兔纤维中张力维持程度存在分歧,我重新研究了甘油提取的兔腰大肌和大田鳖飞行肌在僵直状态以及在MgAMPPNP存在下的力学特性。添加AMPPNP时昆虫纤维会可逆地伸长;该效应只需少量核苷酸。添加AMPPNP时兔纤维仅表现出相对较小的可逆伸长;因此,我关于机械化学平衡的实验证据仅对昆虫肌肉确定。在低肌肉张力下,两种肌肉类型的伸展产生的张力衰减都很小;观察到的应力松弛(“粘弹性”)遵循低阶幂律。在更大张力下,两种肌肉都表现出第二种张力衰减形式(“屈服”),在高张力下占主导,导致由于拉伸在几分钟内几乎完全丧失添加的张力。低于此张力时,等长张力衰减相对较慢,并且相当大的张力会持续过夜。在AMPPNP中屈服在比僵直状态低得多的张力下占主导,但相对于这种变化后的水平,屈服过程的速度似乎没有太大改变;同样,低于一定水平时张力衰减变得非常缓慢。AMPPNP对屈服的影响是可逆的,并且在两种组织中似乎相似。在使用前对AMPPNP进行纯化或添加肌酸激酶抑制剂和ADP去除酶系统时,这些特性均未发生明显变化。因此,在两种制剂中,张力相对于单个横桥的预期寿命维持的时间都很长,并且在屈服过程中其衰减不会因缩短该寿命而加速。这些现象表明,除了横桥分离之外的某些过程正在限制横桥滑动;我正在从相邻横桥之间的相互作用方面寻找解释。