Ion channels: the concept emerges

Angus M. Brown
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Abstract

The belief that the broad sweep of the Hodgkin–Huxley papers (Hodgkin & Huxley, 1952a,c,e,f Hodgkin et al. 1952) can be adequately summarised within a standard textbook chapter is fanciful at best and disingenuous at worst, because such summaries consist of scientific sound-bites that deprive students of the big picture; how decisive experiments create a linear narrative culminating in an internally consistent piece of work. In textbook chapters devoted to Hodgkin and Huxley’s work there are two topics that are inevitably omitted. The first is the separation of the trans-membrane current into INa and IK, the process usually described as a simple subtraction of currents recorded in Na+-free seawater from control currents, which is a gross simplification that neglects to recognise Hodgkin and Huxley’s elegant mathematical solution to the problem (Hodgkin & Huxley, 1952c). Fortunately there exists a superb account of this subject (Cronin, 1987). The second topic, and the subject of this editorial, relates to whether ion movements across a membrane conform to the independence principle. This principle, which was universally accepted at the time, derived from the equations of Teorell and Ussing, and described how the probability of the movement of ions across membranes was under the influence of electrical and chemical gradients but was independent of the presence of other ions (Teorell, 1949; Ussing, 1949). This topic appears deceptively simple upon initial inspection but consideration of the underlying mathematical foundations reveals unexpected complexity. Consider a cell bathed in saline where the influx of a particular ion can be expressed as
离子通道:这个概念出现了
认为霍奇金-赫胥黎论文(Hodgkin & Huxley, 1952a,c,e,f Hodgkin et al. 1952)的广泛内容可以在一个标准的教科书章节中得到充分总结,这种想法充其量是幻想,最坏的情况下是虚伪的,因为这样的总结由科学的声音片段组成,剥夺了学生的大局;决定性的实验如何创造一个线性叙事,最终形成一个内部一致的作品。在教科书中专门介绍霍奇金和赫胥黎作品的章节中,有两个主题不可避免地被省略了。第一种是将跨膜电流分离为INa和IK,这一过程通常被描述为将无Na+海水中记录的电流从控制电流中简单地减去,这是一种粗略的简化,忽略了霍奇金和赫胥黎对该问题的优雅数学解决方案(Hodgkin & Huxley, 1952c)。幸运的是,有一个关于这个主题的极好的描述(Cronin, 1987)。第二个主题,也是这篇社论的主题,涉及到离子在膜上的运动是否符合独立性原则。这一原理在当时被普遍接受,它是从Teorell和Ussing的方程中推导出来的,描述了离子在膜上运动的概率如何受到电和化学梯度的影响,但与其他离子的存在无关(Teorell, 1949;美国,1949年)。这个主题在最初的检查中看似简单,但考虑到潜在的数学基础,就会发现意想不到的复杂性。考虑一个浸泡在盐水中的细胞,其中一个特定离子的流入可以表示为
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