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一些植物（例如捕蝇草（''Dionaea muscipula，也叫 Venus flytrap''））使用钠门控的通道操作运动，本质上是“计数”。捕蝇草可以在北卡罗来纳州和南卡罗来纳州的亚热带湿地见到。<ref name=":16">{{Cite journal|last=Luken|first=James O. | name-list-style = vanc |date= December 2005 |title=Habitats of Dionaea muscipula (Venus' Fly Trap), Droseraceae, Associated with Carolina Bays|journal=Southeastern Naturalist|language=en|volume=4|issue=4|pages=573–584|doi=10.1656/1528-7092(2005)004[0573:HODMVF]2.0.CO;2|issn=1528-7092}}</ref> 当土壤养分不足时，捕蝇草依靠昆虫和动物为食。<ref name=":1">{{cite journal | vauthors = Böhm J, Scherzer S, Krol E, Kreuzer I, von Meyer K, Lorey C, Mueller TD, Shabala L, Monte I, Solano R, Al-Rasheid KA, Rennenberg H, Shabala S, Neher E, Hedrich R | display-authors = 6 | title = The Venus Flytrap Dionaea muscipula Counts Prey-Induced Action Potentials to Induce Sodium Uptake | journal = Current Biology | volume = 26 | issue = 3 | pages = 286–95 | date = February 2016 | pmid = 26804557 | pmc = 4751343 | doi = 10.1016/j.cub.2015.11.057 }}</ref> 尽管对这种植物进行了研究，但对于捕蝇草以及一般的食肉植物的分子机制还缺乏了解。<ref name=":2">{{cite journal | vauthors = Hedrich R, Neher E | title = Venus Flytrap: How an Excitable, Carnivorous Plant Works | journal = Trends in Plant Science | volume = 23 | issue = 3 | pages = 220–234 | date = March 2018 | pmid = 29336976 | doi = 10.1016/j.tplants.2017.12.004 }}</ref>

一些植物（例如捕蝇草（''Dionaea muscipula，也叫 Venus flytrap''））使用钠门控的通道操作运动，本质上是“计数”。捕蝇草可以在北卡罗来纳州和南卡罗来纳州的亚热带湿地见到。<ref name=":16">{{Cite journal|last=Luken|first=James O. | name-list-style = vanc |date= December 2005 |title=Habitats of Dionaea muscipula (Venus' Fly Trap), Droseraceae, Associated with Carolina Bays|journal=Southeastern Naturalist|language=en|volume=4|issue=4|pages=573–584|doi=10.1656/1528-7092(2005)004[0573:HODMVF]2.0.CO;2|issn=1528-7092}}</ref> 当土壤养分不足时，捕蝇草依靠昆虫和动物为食。<ref name=":1">{{cite journal | vauthors = Böhm J, Scherzer S, Krol E, Kreuzer I, von Meyer K, Lorey C, Mueller TD, Shabala L, Monte I, Solano R, Al-Rasheid KA, Rennenberg H, Shabala S, Neher E, Hedrich R | display-authors = 6 | title = The Venus Flytrap Dionaea muscipula Counts Prey-Induced Action Potentials to Induce Sodium Uptake | journal = Current Biology | volume = 26 | issue = 3 | pages = 286–95 | date = February 2016 | pmid = 26804557 | pmc = 4751343 | doi = 10.1016/j.cub.2015.11.057 }}</ref> 尽管对这种植物进行了研究，但对于捕蝇草以及一般的食肉植物的分子机制还缺乏了解。<ref name=":2">{{cite journal | vauthors = Hedrich R, Neher E | title = Venus Flytrap: How an Excitable, Carnivorous Plant Works | journal = Trends in Plant Science | volume = 23 | issue = 3 | pages = 220–234 | date = March 2018 | pmid = 29336976 | doi = 10.1016/j.tplants.2017.12.004 }}</ref>

不过，已经有很多关于捕蝇草的动作电位及其对捕蝇草的运动和机械原理的作用研究。首先，捕蝇草的静息膜电位（-120 mV）低于动物细胞（通常为 -90 mV 至 -40 mV）。<ref name=":2" /><ref name=":17">Purves D, Augustine GJ, Fitzpatrick D, et al., editors. Neuroscience. 2nd edition. Sunderland (MA): Sinauer Associates; 2001. Electrical Potentials Across Nerve Cell Membranes.Available from: <nowiki>https://www.ncbi.nlm.nih.gov/books/NBK11069/</nowiki></ref> 更低的静息电位使得动作电位更容易激发。因此，当一只昆虫落在植物的陷阱上时，它就会触发一个毛发样的机械感受器。<ref name=":2" /> 这个受体激活一个持续约 1.5 毫秒的动作电位。<ref name=":18">{{cite journal | vauthors = Volkov AG, Adesina T, Jovanov E | title = Closing of venus flytrap by electrical stimulation of motor cells | journal = Plant Signaling & Behavior | volume = 2 | issue = 3 | pages = 139–45 | date = May 2007 | pmid = 19516982 | pmc = 2634039 | doi = 10.4161/psb.2.3.4217 }}</ref> 最终，这会导致钙离子进入细胞，使细胞稍微去极化。

不过，已经有很多关于捕蝇草的动作电位及其对捕蝇草的运动和机械原理的作用研究。首先，捕蝇草的静息膜电位（-120 mV）低于动物细胞（通常为 -90 mV 至 -40 mV）。<ref name=":2" /><ref name=":17">Purves D, Augustine GJ, Fitzpatrick D, et al., editors. Neuroscience. 2nd edition. Sunderland (MA): Sinauer Associates; 2001. Electrical Potentials Across Nerve Cell Membranes.Available from: <nowiki>https://www.ncbi.nlm.nih.gov/books/NBK11069/</nowiki></ref> 更低的静息电位使得动作电位更容易触发。因此，当昆虫落在植物的陷阱叶上时，会触发一个毛状的机械感受器。<ref name=":2" /> 这个受体激活一个持续约 1.5 毫秒的动作电位。<ref name=":18">{{cite journal | vauthors = Volkov AG, Adesina T, Jovanov E | title = Closing of venus flytrap by electrical stimulation of motor cells | journal = Plant Signaling & Behavior | volume = 2 | issue = 3 | pages = 139–45 | date = May 2007 | pmid = 19516982 | pmc = 2634039 | doi = 10.4161/psb.2.3.4217 }}</ref> 最终，这会导致钙离子进入细胞，使细胞稍微去极化。

However, the flytrap doesn't close after one trigger. Instead, it requires the activation of 2 or more hairs.<ref name=":1" /><ref name=":2" /> If only one hair is triggered, it throws the activation as a false positive. Further, the second hair must be activated within a certain time interval （0.75 s - 40 s） for it to register with the first activation.<ref name=":2" /> Thus, a buildup of calcium starts and slowly falls from the first trigger. When the second action potential is fired within the time interval, it reaches the Calcium threshold to depolarize the cell, closing the trap on the prey within a fraction of a second.<ref name=":2" />

然而，一次触发不会让捕蝇草闭合，而是需要 2 根或更多的感觉毛被激活。<ref name=":1" /><ref name=":2" /> 只有一根感觉毛被触发，这个激活会被视为假阳性而无反应。而且第二根毛必须在一定的时间间隔（0.75 s - 40 s）内被激活，才能与前次激活一起算。<ref name=":2" /> 因此，一次触发让钙信号开始积累并缓慢下降，而第二个动作电位在特定时间内发放，让钙信号达到钙阈值而使细胞去极化，并在一秒内闭合陷阱叶。<ref name=":2" />

与随后的阳性的钾离子释放一样，植物的动作电位与盐（KCl）的渗透流失（osmotic loss）有关。然而，动物的动作电位是渗透中性的，因为等量的钠内流和钾外流在渗透压上相互抵消。植物细胞<ref name="Gradmann_1998" group="lower-alpha">{{cite journal | vauthors = Gradmann D, Hoffstadt J | title = Electrocoupling of ion transporters in plants: interaction with internal ion concentrations | journal = The Journal of Membrane Biology | volume = 166 | issue = 1 | pages = 51–9 | date = November 1998 | pmid = 9784585 | doi = 10.1007/s002329900446 | s2cid = 24190001 }}</ref>中电和渗透关系的相互作用似乎起源于盐度（salinity）变化条件下动植物共同的单细胞祖先的电兴奋性的渗透作用。此外，现在的快速信号传递功能认为是后生动物（metazoan）细胞在更稳定的渗透环境中的更新的发展。<ref name="Gradmann_1980">

 

Gradmann, D; Mummert, H in {{harvnb|Spanswick|Lucas|Dainty|1980|loc=''Plant action potentials'', pp. 333–344.}}</ref> 动作电位在一些维管植物比如含羞草（''Mimosa pudica''）中我们熟悉的信号传递功能，可能是独立于后生动物兴奋性细胞而产生的。

Together with the subsequent release of positive potassium ions the action potential in plants involves an [[osmotic]] loss of salt （KCl）. Whereas, the animal action potential is osmotically neutral because equal amounts of entering sodium and leaving potassium cancel each other osmotically. The interaction of electrical and osmotic relations in plant cells<ref name="Gradmann_1998" group="lower-alpha">{{cite journal | vauthors = Gradmann D, Hoffstadt J | title = Electrocoupling of ion transporters in plants: interaction with internal ion concentrations | journal = The Journal of Membrane Biology | volume = 166 | issue = 1 | pages = 51–9 | date = November 1998 | pmid = 9784585 | doi = 10.1007/s002329900446 | s2cid = 24190001 }}</ref> appears to have arisen from an osmotic function of electrical excitability in a common unicellular ancestors of plants and animals under changing salinity conditions. Further, the present function of rapid signal transmission is seen as a newer accomplishment of [[metazoan]] cells in a more stable osmotic environment.<ref name="Gradmann_1980">

Gradmann, D; Mummert, H in {{harvnb|Spanswick|Lucas|Dainty|1980|loc=''Plant action potentials'', pp. 333–344.}}</ref> It is likely that the familiar signaling function of action potentials in some vascular plants （e.g. ''[[Mimosa pudica]]''） arose independently from that in metazoan excitable cells.

 

不同于上升相和峰值，下降相和后超极化似乎主要依赖于非钙离子的阳离子。为了启动复极化，细胞需要将钾离子通过细胞膜上的被动运输机制输出到胞外。不像神经元中钾离子的移动决定膜电位下降，植物细胞完全复极化时，需要消耗 ATP 能，帮助细胞利用称为氢钾 ATP 酶（ H⁺-ATPase）的转运蛋白释放氢。<ref name="Opritov">Opritov, V A, et al. “Direct Coupling of Action Potential Generation in Cells of a Higher Plant (Cucurbita Pepo) with the Operation of an Electrogenic Pump.” ''Russian Journal of Plant Physiology'', vol. 49, no. 1, 2002, pp. 142–147.</ref><ref name=":2" />

不同于上升相和峰值，下降相和后超极化似乎主要依赖于非钙离子的阳离子。为了启动复极化，细胞需要将钾离子通过细胞膜上的被动运输机制输出到胞外。不像神经元中钾离子的移动决定膜电位下降，植物细胞完全复极化时，需要消耗 ATP 能，帮助细胞利用称为氢钾 ATP 酶（ H⁺-ATPase）的转运蛋白释放氢。<ref name="Opritov">Opritov, V A, et al. “Direct Coupling of Action Potential Generation in Cells of a Higher Plant (Cucurbita Pepo) with the Operation of an Electrogenic Pump.” ''Russian Journal of Plant Physiology'', vol. 49, no. 1, 2002, pp. 142–147.</ref><ref name=":2" />

第一个问题通过研究枪乌贼（''[[Loligo forbesii]]'' 和 ''[[Doryteuthis pealeii]]'', 当时被归类为 ''Loligo pealeii''）神经元中发现的巨大轴突得以解决。<ref name="keynes_1989" group="lower-alpha">{{cite journal | vauthors = Keynes RD | title = The role of giant axons in studies of the nerve impulse | journal = BioEssays | volume = 10 | issue = 2–3 | pages = 90–3 | year = 1989 | pmid = 2541698 | doi = 10.1002/bies.950100213 }}</ref> 这些轴突直径很大（约 1 毫米，比一个典型的神经元大 100 倍），肉眼可见，因此很容易提取和操作。<ref name="hodgkin_1952" group="lower-alpha" /><ref name="Meunier" group="lower-alpha">{{cite journal | vauthors = Meunier C, Segev I | title = Playing the devil's advocate: is the Hodgkin-Huxley model useful? | journal = Trends in Neurosciences | volume = 25 | issue = 11 | pages = 558–63 | date = November 2002 | pmid = 12392930 | doi = 10.1016/S0166-2236(02)02278-6 | s2cid = 1355280 }}</ref> 不过，它们并不代表所有的兴奋性细胞，人们也研究了多种其他的动作电位系统。

第一个问题通过研究枪乌贼（''[[Loligo forbesii]]'' 和 ''[[Doryteuthis pealeii]]'', 当时被归类为 ''Loligo pealeii''）神经元中发现的巨大轴突得以解决。<ref name="keynes_1989" group="lower-alpha">{{cite journal | vauthors = Keynes RD | title = The role of giant axons in studies of the nerve impulse | journal = BioEssays | volume = 10 | issue = 2–3 | pages = 90–3 | year = 1989 | pmid = 2541698 | doi = 10.1002/bies.950100213 }}</ref> 这些轴突直径很大（约 1 毫米，比一个典型的神经元大 100 倍），肉眼可见，因此很容易提取和操作。<ref name="hodgkin_1952" group="lower-alpha" /><ref name="Meunier" group="lower-alpha">{{cite journal | vauthors = Meunier C, Segev I | title = Playing the devil's advocate: is the Hodgkin-Huxley model useful? | journal = Trends in Neurosciences | volume = 25 | issue = 11 | pages = 558–63 | date = November 2002 | pmid = 12392930 | doi = 10.1016/S0166-2236(02)02278-6 | s2cid = 1355280 }}</ref> 不过，它们并不代表所有的兴奋性细胞，人们也研究了多种其他的动作电位系统。

第二个问题随着电压钳<ref name="cole_1949" group="lower-alpha">{{cite journal | vauthors = Cole KS | year = 1949 | title = Dynamic electrical characteristics of the squid axon membrane | journal = Arch. Sci. Physiol. | volume = 3 | pages = 253–8| author-link = Kenneth Stewart Cole }}</ref> 关键性的开发得以解决，让实验者可以对动作电位的离子电流进行单独研究，并消除了电子噪声的主要来源——与膜电容 ''C'' 关联的电流 ''I_C''。因为电流等于''C'' 乘以跨膜电压 ''V_m'' 的变化率，所以解决方案就是设计一个电路，让 ''V_m'' 保持固定（零变化率），不管跨膜电流的变化。因此，使 ''V_m'' 保持固定所需的电流是流过膜的电流直接取反。电子设备的其他进步包括法拉第笼（Faraday cages）和具有高输入阻抗的电子器件的使用，这样测量本身就不会影响被测量的电位。

第二个问题随着电压钳（voltage clamp）<ref name="cole_1949" group="lower-alpha">{{cite journal | vauthors = Cole KS | year = 1949 | title = Dynamic electrical characteristics of the squid axon membrane | journal = Arch. Sci. Physiol. | volume = 3 | pages = 253–8| author-link = Kenneth Stewart Cole }}</ref> 关键性的开发得以解决，让实验者可以对动作电位的离子电流进行单独研究，并消除了电子噪声的主要来源——与膜电容 ''C'' 关联的电流 ''I_C''。因为电流等于''C'' 乘以跨膜电压 ''V_m'' 的变化率，所以解决方案就是设计一个电路，让 ''V_m'' 保持固定（零变化率），不管跨膜电流的变化。因此，使 ''V_m'' 保持固定所需的电流是流过膜的电流直接取反。电子设备的其他进步包括法拉第笼（Faraday cages）和具有高输入阻抗的电子器件的使用，这样测量本身就不会影响被测量的电位。

第三个问题是如何获得足够小的电极来记录单个轴突内的电位而不对其造成干扰，这个问题在 1949 年因玻璃微电极的发明而得以解决。<ref name="ling_1949" group="lower-alpha">{{cite journal | vauthors = Ling G, Gerard RW | title = The normal membrane potential of frog sartorius fibers | journal = Journal of Cellular and Comparative Physiology | volume = 34 | issue = 3 | pages = 383–96 | date = December 1949 | pmid = 15410483 | doi = 10.1002/jcp.1030340304 }}</ref> 玻璃电极很快被其他研究人员所采用。<ref name="nastuk_1950" group="lower-alpha">{{cite journal | vauthors = Nastuk WL, Hodgkin A | year = 1950 | title = The electrical activity of single muscle fibers | journal = Journal of Cellular and Comparative Physiology | volume = 35 | pages = 39–73 | doi = 10.1002/jcp.1030350105 }}</ref><ref name="brock_1952" group="lower-alpha">{{cite journal | vauthors = Brock LG, Coombs JS, Eccles JC | title = The recording of potentials from motoneurones with an intracellular electrode | journal = The Journal of Physiology | volume = 117 | issue = 4 | pages = 431–60 | date = August 1952 | pmid = 12991232 | pmc = 1392415 | doi = 10.1113/jphysiol.1952.sp004759 }}</ref> 这种方法改进后可以生产出细到 100 Å（10 nm）且有高输入阻抗的电极端部。<ref name=":20">Snell, FM in {{harvnb|Lavallée|Schanne|Hébert|1969|loc=''Some Electrical Properties of Fine-Tipped Pipette Microelectrodes''.}}</ref> 动作电位也可用置于神经元附近的小金属电极、或电解质氧化物半导体场效应晶体管（EOSFET）的神经芯片（neurochip）进行记录，或利用 Ca²⁺ 或电压敏感的染料进行光学记录。<ref name="dyes" group="lower-alpha">{{cite journal | vauthors = Ross WN, Salzberg BM, Cohen LB, Davila HV | title = A large change in dye absorption during the action potential | journal = Biophysical Journal | volume = 14 | issue = 12 | pages = 983–6 | date = December 1974 | pmid = 4429774 | pmc = 1334592 | doi = 10.1016/S0006-3495(74)85963-1 | bibcode = 1974BpJ....14..983R }}<br />* {{cite journal | vauthors = Grynkiewicz G, Poenie M, Tsien RY | title = A new generation of Ca2+ indicators with greatly improved fluorescence properties | journal = The Journal of Biological Chemistry | volume = 260 | issue = 6 | pages = 3440–50 | date = March 1985 | doi = 10.1016/S0021-9258(19)83641-4 | pmid = 3838314 | doi-access = free }}</ref>[[Image:Single channel.png|thumb|left|电压钳电极揭示了，离子通道有两个状态：开放（高电导）和关闭（低电导）。|链接=Special:FilePath/Single_channel.png]]

第三个问题是如何获得足够小的电极来记录单个轴突内的电位而不对其造成干扰，这个问题在 1949 年因玻璃微电极的发明而得以解决。<ref name="ling_1949" group="lower-alpha">{{cite journal | vauthors = Ling G, Gerard RW | title = The normal membrane potential of frog sartorius fibers | journal = Journal of Cellular and Comparative Physiology | volume = 34 | issue = 3 | pages = 383–96 | date = December 1949 | pmid = 15410483 | doi = 10.1002/jcp.1030340304 }}</ref> 玻璃电极很快被其他研究人员所采用。<ref name="nastuk_1950" group="lower-alpha">{{cite journal | vauthors = Nastuk WL, Hodgkin A | year = 1950 | title = The electrical activity of single muscle fibers | journal = Journal of Cellular and Comparative Physiology | volume = 35 | pages = 39–73 | doi = 10.1002/jcp.1030350105 }}</ref><ref name="brock_1952" group="lower-alpha">{{cite journal | vauthors = Brock LG, Coombs JS, Eccles JC | title = The recording of potentials from motoneurones with an intracellular electrode | journal = The Journal of Physiology | volume = 117 | issue = 4 | pages = 431–60 | date = August 1952 | pmid = 12991232 | pmc = 1392415 | doi = 10.1113/jphysiol.1952.sp004759 }}</ref> 这种方法改进后可以生产出细到 100 Å（10 nm）且有高输入阻抗的电极端部。<ref name=":20">Snell, FM in {{harvnb|Lavallée|Schanne|Hébert|1969|loc=''Some Electrical Properties of Fine-Tipped Pipette Microelectrodes''.}}</ref> 动作电位也可用置于神经元附近的小金属电极、或电解质氧化物半导体场效应晶体管（EOSFET）的神经芯片（neurochip）进行记录，或利用 Ca²⁺ 或电压敏感的染料进行光学记录。<ref name="dyes" group="lower-alpha">{{cite journal | vauthors = Ross WN, Salzberg BM, Cohen LB, Davila HV | title = A large change in dye absorption during the action potential | journal = Biophysical Journal | volume = 14 | issue = 12 | pages = 983–6 | date = December 1974 | pmid = 4429774 | pmc = 1334592 | doi = 10.1016/S0006-3495(74)85963-1 | bibcode = 1974BpJ....14..983R }}<br />* {{cite journal | vauthors = Grynkiewicz G, Poenie M, Tsien RY | title = A new generation of Ca2+ indicators with greatly improved fluorescence properties | journal = The Journal of Biological Chemistry | volume = 260 | issue = 6 | pages = 3440–50 | date = March 1985 | doi = 10.1016/S0021-9258(19)83641-4 | pmid = 3838314 | doi-access = free }}</ref>[[Image:Single channel.png|thumb|left|电压钳电极揭示了，离子通道有两个状态：开放（高电导）和关闭（低电导）。|链接=Special:FilePath/Single_channel.png]]

电在动物神经系统中的作用最早是由 Luigi Galvani 在解剖的青蛙中观察到的，他在 1791 年到 1797 年研究了这一现象。Galvani 的研究结果促成Alessandro Volta 发明了伏打电堆（voltaic pile）——已知最早的电池——他用这种电池研究了动物电（如电鳗）以及对直流电压的生理反应。<ref name="piccolino_2000" group="lower-alpha">{{cite journal | vauthors = Piccolino M | title = The bicentennial of the Voltaic battery (1800-2000): the artificial electric organ | journal = Trends in Neurosciences | volume = 23 | issue = 4 | pages = 147–51 | date = April 2000 | pmid = 10717671 | doi = 10.1016/S0166-2236(99)01544-1 | s2cid = 393323 }}</ref>

电在动物神经系统中的作用最早是由 Luigi Galvani 在解剖的青蛙中观察到的，他在 1791 年到 1797 年研究了这一现象。Galvani 的研究结果促成Alessandro Volta 发明了伏打电堆（voltaic pile）——已知最早的电池——他用这种电池研究了动物电（如电鳗）以及对直流电压的生理反应。<ref name="piccolino_2000" group="lower-alpha">{{cite journal | vauthors = Piccolino M | title = The bicentennial of the Voltaic battery (1800-2000): the artificial electric organ | journal = Trends in Neurosciences | volume = 23 | issue = 4 | pages = 147–51 | date = April 2000 | pmid = 10717671 | doi = 10.1016/S0166-2236(99)01544-1 | s2cid = 393323 }}</ref>

Scientists of the 19th century studied the propagation of electrical signals in whole [[nerve]]s （i.e., bundles of [[neuron]]s） and demonstrated that nervous tissue was made up of [[cell (biology)|cells]], instead of an interconnected network of tubes （a ''reticulum''）.{{sfnm|1a1=Brazier|1y=1961|2a1=McHenry|2a2=Garrison|2y=1969|3a1=Worden|3a2=Swazey|3a3=Adelman|3y=1975}} [[Carlo Matteucci]] followed up Galvani's studies and demonstrated that [[cell membrane]]s had a voltage across them and could produce [[direct current]]. Matteucci's work inspired the German physiologist, [[Emil du Bois-Reymond]], who discovered the action potential in 1843.<ref name=":21">{{Cite book|title=Emil du Bois-Reymond : neuroscience, self, and society in nineteenth-century Germany|last=Finkelstein | first = Gabriel Ward | name-list-style = vanc |isbn=9781461950325|location=Cambridge, Massachusetts|oclc=864592470|year = 2013}}</ref> The [[conduction velocity]] of action potentials was first measured in 1850 by du Bois-Reymond's friend, [[Hermann von Helmholtz]].<ref name=":22">[[Kathryn Olesko|Olesko, Kathryn M.]], and Frederic L. Holmes. "Experiment, Quantification and Discovery: Helmholtz's Early Physiological Researches, 1843-50". In ''Hermann von Helmholtz and the Foundations of Nineteenth Century Science'', ed. David Cahan, 50-108. Berkeley; Los Angeles; London: University of California, 1994.</ref> To establish that nervous tissue is made up of discrete cells, the Spanish physician [[Santiago Ramón y Cajal]] and his students used a stain developed by [[Camillo Golgi]] to reveal the myriad shapes of neurons, which they rendered painstakingly. For their discoveries, Golgi and Ramón y Cajal were awarded the 1906 [[Nobel Prize in Physiology or Medicine|Nobel Prize in Physiology]].<ref name="Nobel_1906" group="lower-Greek">{{cite press release | url = http://nobelprize.org/medicine/laureates/1906/index.html | title = The Nobel Prize in Physiology or Medicine 1906 | publisher = The Royal Swedish Academy of Science | year = 1906 | access-date = 2010-02-21 | url-status = live | archive-url = https://web.archive.org/web/20081204190959/http://nobelprize.org/medicine/laureates/1906/index.html | archive-date = 4 December 2008 | df = dmy-all }}</ref> Their work resolved a long-standing controversy in the [[neuroanatomy]] of the 19th century; Golgi himself had argued for the network model of the nervous system.

19 世纪的科学家研究了电信号在整个神经（nerves）（即神经元束）中的传播，并证明神经组织是由细胞组成的，而不是一个互相连接的管网（''reticulum''）。Carlo Matteucci 继续 Galvani 的研究，证明细胞膜上存在电压，可以产生直流电。受 Carlo Matteucci 的工作启发，德国生理学家 [[Emil du Bois-Reymond]] 在1843 年发现了动作电位。<ref name=":21">{{Cite book|title=Emil du Bois-Reymond : neuroscience, self, and society in nineteenth-century Germany|last=Finkelstein | first = Gabriel Ward | name-list-style = vanc |isbn=9781461950325|location=Cambridge, Massachusetts|oclc=864592470|year = 2013}}</ref> 动作电位的传导速率最早是在 1850 年由 Hermann von Helmholtz 测量的。<ref name=":22">[[Kathryn Olesko|Olesko, Kathryn M.]], and Frederic L. Holmes. "Experiment, Quantification and Discovery: Helmholtz's Early Physiological Researches, 1843-50". In ''Hermann von Helmholtz and the Foundations of Nineteenth Century Science'', ed. David Cahan, 50-108. Berkeley; Los Angeles; London: University of California, 1994.</ref> 为了证明神经组织是由独立的细胞组成的，西班牙物理学家 [[Santiago Ramón y Cajal]] 和他的学生们使用 Camillo Golgi 开发的染色剂揭示了大量的神经元形状，他们煞费苦心地进行了绘制。由于他们的发现，Golgi 和 [[Santiago Ramón y Cajal|Ramón y Cajal]] 获得了 1906 年的诺贝尔生理学奖。<ref name="Nobel_1906" group="lower-Greek">{{cite press release | url = http://nobelprize.org/medicine/laureates/1906/index.html | title = The Nobel Prize in Physiology or Medicine 1906 | publisher = The Royal Swedish Academy of Science | year = 1906 | access-date = 2010-02-21 | url-status = live | archive-url = https://web.archive.org/web/20081204190959/http://nobelprize.org/medicine/laureates/1906/index.html | archive-date = 4 December 2008 | df = dmy-all }}</ref> 他们的工作解决了 19 世纪神经解剖学中长期存在的争议；高尔基自己则坚持神经系统的管网模型。

 

19 世纪的科学家研究了电信号在整个神经（nerves）（即神经元束）中的传播，并证明神经组织是由细胞组成的，而不是一个互相连接的管网（''reticulum''）。Carlo Matteucci 继续 Galvani 的研究，证明细胞膜上存在电压，可以产生直流电。受 Carlo Matteucci 的工作启发，德国生理学家 [[Emil du Bois-Reymond]] 在1843 年发现了动作电位。<ref name=":21" /> 动作电位的传导速率最早是在 1850 年由 Hermann von Helmholtz 测量的。<ref name=":22" /> 为了证明神经组织是由独立的细胞组成的，西班牙物理学家 [[Santiago Ramón y Cajal]] 和他的学生们使用 Camillo Golgi 开发的染色剂揭示了大量的神经元形状，他们煞费苦心地进行了绘制。由于他们的发现，Golgi 和 [[Santiago Ramón y Cajal|Ramón y Cajal]] 获得了 1906 年的诺贝尔生理学奖。<ref name="Nobel_1906" group="lower-Greek" /> 他们的工作解决了 19 世纪神经解剖学中长期存在的争议；高尔基自己则坚持神经系统的网络模型。

[[Image:3b8e.png|thumb|right|钠钾泵在 E2-Pi 状态下的带状图（Ribbon diagram）。推测的脂双层边界显示为蓝色（胞内）和红色（胞外）的面。|链接=Special:FilePath/3b8e.png]]

[[Image:3b8e.png|thumb|right|钠钾泵在 E2-Pi 状态下的带状图（Ribbon diagram）。推测的脂双层边界显示为蓝色（胞内）和红色（胞外）的面。|链接=Special:FilePath/3b8e.png]]

The 20th century was a significant era for electrophysiology. In 1902 and again in 1912, [[Julius Bernstein]] advanced the hypothesis that the action potential resulted from a change in the [[permeation|permeability]] of the axonal membrane to ions.<ref name="bernstein_1902_1912" group="lower-alpha">{{cite journal | vauthors = Bernstein J | year = 1902 | title = Untersuchungen zur Thermodynamik der bioelektrischen Ströme | journal = Pflügers Archiv für die gesamte Physiologie | volume = 92 | pages = 521–562 | doi = 10.1007/BF01790181 | issue = 10–12| s2cid = 33229139 | author-link = Julius Bernstein | url = https://zenodo.org/record/2192363 }}</ref>{{sfn|Bernstein|1912}} Bernstein's hypothesis was confirmed by [[Kenneth Stewart Cole|Ken Cole]] and Howard Curtis, who showed that membrane conductance increases during an action potential.<ref group="lower-alpha" name=":16">{{cite journal | vauthors = Cole KS, Curtis HJ | title = Electric Impedance of the Squid Giant Axon During Activity | journal = The Journal of General Physiology | volume = 22 | issue = 5 | pages = 649–70 | date = May 1939 | pmid = 19873125 | pmc = 2142006 | doi = 10.1085/jgp.22.5.649 | author-link1 = Kenneth Stewart Cole }}</ref> In 1907, [[Louis Lapicque]] suggested that the action potential was generated as a threshold was crossed,<ref group="lower-alpha" name=":17">{{cite journal | vauthors = Lapicque L | year = 1907 | title = Recherches quantitatives sur l'excitationelectrique des nerfs traitee comme une polarisation | journal = J. Physiol. Pathol. Gen | volume = 9| pages = 620–635 }}</ref> what would be later shown as a product of the [[dynamical system]]s of ionic conductances. In 1949, [[Alan Lloyd Hodgkin|Alan Hodgkin]] and [[Bernard Katz]] refined Bernstein's hypothesis by considering that the axonal membrane might have different permeabilities to different ions; in particular, they demonstrated the crucial role of the sodium permeability for the action potential.<ref name="hodgkin_1949" group="lower-alpha">{{cite journal | vauthors = Hodgkin AL, Katz B | title = The effect of sodium ions on the electrical activity of giant axon of the squid | journal = The Journal of Physiology | volume = 108 | issue = 1 | pages = 37–77 | date = March 1949 | pmid = 18128147 | pmc = 1392331 | doi = 10.1113/jphysiol.1949.sp004310 | author-link1 = Alan Lloyd Hodgkin | author-link2 = Bernard Katz }}</ref> They made the first actual recording of the electrical changes across the neuronal membrane that mediate the action potential.<ref group="lower-Greek" name=":0">{{cite journal |last=Warlow|first=Charles| name-list-style = vanc |title=The Recent Evolution of a Symbiotic Ion Channel in the Legume Family Altered Ion Conductance and Improved Functionality in Calcium Signaling|journal=Practical Neurology|volume=7|issue=3|pages=192–197|url=http://pn.bmj.com/content/7/3/192.full|publisher=BMJ Publishing Group|access-date=23 March 2013|url-status=live|archive-url=https://web.archive.org/web/20120314104408/http://pn.bmj.com/content/7/3/192.full|archive-date=14 March 2012|df=dmy-all|date=June 2007}}</ref> This line of research culminated in the five 1952 papers of Hodgkin, Katz and [[Andrew Huxley]], in which they applied the [[voltage clamp]] technique to determine the dependence of the axonal membrane's permeabilities to sodium and potassium ions on voltage and time, from which they were able to reconstruct the action potential quantitatively.<ref name="hodgkin_1952" group="lower-alpha" /> Hodgkin and Huxley correlated the properties of their mathematical model with discrete [[ion channel]]s that could exist in several different states, including "open", "closed", and "inactivated". Their hypotheses were confirmed in the mid-1970s and 1980s by [[Erwin Neher]] and [[Bert Sakmann]], who developed the technique of [[patch clamp]]ing to examine the conductance states of individual ion channels.<ref name="patch_clamp" group="lower-alpha">{{cite journal | vauthors = Neher E, Sakmann B | title = Single-channel currents recorded from membrane of denervated frog muscle fibres | journal = Nature | volume = 260 | issue = 5554 | pages = 799–802 | date = April 1976 | pmid = 1083489 | doi = 10.1038/260799a0 | author-link1 = Erwin Neher | bibcode = 1976Natur.260..799N | s2cid = 4204985 }}<br />* {{cite journal | vauthors = Hamill OP, Marty A, Neher E, Sakmann B, Sigworth FJ | title = Improved patch-clamp techniques for high-resolution current recording from cells and cell-free membrane patches | journal = Pflügers Archiv | volume = 391 | issue = 2 | pages = 85–100 | date = August 1981 | pmid = 6270629 | doi = 10.1007/BF00656997 | s2cid = 12014433 }}<br />* {{cite journal | vauthors = Neher E, Sakmann B | title = The patch clamp technique | journal = Scientific American | volume = 266 | issue = 3 | pages = 44–51 | date = March 1992 | pmid = 1374932 | doi = 10.1038/scientificamerican0392-44 | author-link1 = Erwin Neher | bibcode = 1992SciAm.266c..44N }}</ref> In the 21st century, researchers are beginning to understand the structural basis for these conductance states and for the selectivity of channels for their species of ion,<ref name="yellen_2002" group="lower-alpha">{{cite journal | vauthors = Yellen G | title = The voltage-gated potassium channels and their relatives | journal = Nature | volume = 419 | issue = 6902 | pages = 35–42 | date = September 2002 | pmid = 12214225 | doi = 10.1038/nature00978 | bibcode = 2002Natur.419...35Y | s2cid = 4420877 }}</ref> through the atomic-resolution [[X-ray crystallography|crystal structures]],<ref name="doyle_1998" group="lower-alpha">{{cite journal | vauthors = Doyle DA, Morais Cabral J, Pfuetzner RA, Kuo A, Gulbis JM, Cohen SL, Chait BT, MacKinnon R | display-authors = 6 | title = The structure of the potassium channel: molecular basis of K+ conduction and selectivity | journal = Science | volume = 280 | issue = 5360 | pages = 69–77 | date = April 1998 | pmid = 9525859 | doi = 10.1126/science.280.5360.69 | bibcode = 1998Sci...280...69D }}<br />* {{cite journal | vauthors = Zhou Y, Morais-Cabral JH, Kaufman A, MacKinnon R | title = Chemistry of ion coordination and hydration revealed by a K+ channel-Fab complex at 2.0 A resolution | journal = Nature | volume = 414 | issue = 6859 | pages = 43–8 | date = November 2001 | pmid = 11689936 | doi = 10.1038/35102009 | bibcode = 2001Natur.414...43Z | s2cid = 205022645 }}<br />* {{cite journal | vauthors = Jiang Y, Lee A, Chen J, Ruta V, Cadene M, Chait BT, MacKinnon R | title = X-ray structure of a voltage-dependent K+ channel | journal = Nature | volume = 423 | issue = 6935 | pages = 33–41 | date = May 2003 | pmid = 12721618 | doi = 10.1038/nature01580 | bibcode = 2003Natur.423...33J | s2cid = 4347957 }}</ref> fluorescence distance measurements<ref name="FRET" group="lower-alpha">{{cite journal | vauthors = Cha A, Snyder GE, Selvin PR, Bezanilla F | title = Atomic scale movement of the voltage-sensing region in a potassium channel measured via spectroscopy | journal = Nature | volume = 402 | issue = 6763 | pages = 809–13 | date = December 1999 | pmid = 10617201 | doi = 10.1038/45552 | bibcode = 1999Natur.402..809C | s2cid = 4353978 }}<br />* {{cite journal | vauthors = Glauner KS, Mannuzzu LM, Gandhi CS, Isacoff EY | title = Spectroscopic mapping of voltage sensor movement in the Shaker potassium channel | journal = Nature | volume = 402 | issue = 6763 | pages = 813–7 | date = December 1999 | pmid = 10617202 | doi = 10.1038/45561 | bibcode = 1999Natur.402..813G | s2cid = 4417476 }}<br />* {{cite journal | vauthors = Bezanilla F | title = The voltage sensor in voltage-dependent ion channels | journal = Physiological Reviews | volume = 80 | issue = 2 | pages = 555–92 | date = April 2000 | pmid = 10747201 | doi = 10.1152/physrev.2000.80.2.555 }}</ref> and [[cryo-electron microscopy]] studies.<ref name="cryoEM" group="lower-alpha">{{cite journal | vauthors = Catterall WA | title = A 3D view of sodium channels | journal = Nature | volume = 409 | issue = 6823 | pages = 988–9, 991 | date = February 2001 | pmid = 11234048 | doi = 10.1038/35059188 | bibcode = 2001Natur.409..988C | s2cid = 4371677 | doi-access = free }}<br />* {{cite journal | vauthors = Sato C, Ueno Y, Asai K, Takahashi K, Sato M, Engel A, Fujiyoshi Y | title = The voltage-sensitive sodium channel is a bell-shaped molecule with several cavities | journal = Nature | volume = 409 | issue = 6823 | pages = 1047–51 | date = February 2001 | pmid = 11234014 | doi = 10.1038/35059098 | bibcode = 2001Natur.409.1047S | s2cid = 4430165 }}</ref>

20 世纪是电生理的重要时期。Julius Bernstein 于 1902 和 1912 两次提出动作电位是由轴突膜对离子的渗透性的改变引起的假说。<ref name="bernstein_1902_1912" group="lower-alpha">{{cite journal | vauthors = Bernstein J | year = 1902 | title = Untersuchungen zur Thermodynamik der bioelektrischen Ströme | journal = Pflügers Archiv für die gesamte Physiologie | volume = 92 | pages = 521–562 | doi = 10.1007/BF01790181 | issue = 10–12| s2cid = 33229139 | author-link = Julius Bernstein | url = https://zenodo.org/record/2192363 }}</ref> Ken Cole 和 Howard Curtis 证实了 Bernstein 的假说，他们发现在动作电位期间膜电导增加。<ref name=":16" group="lower-alpha">{{cite journal | vauthors = Cole KS, Curtis HJ | title = Electric Impedance of the Squid Giant Axon During Activity | journal = The Journal of General Physiology | volume = 22 | issue = 5 | pages = 649–70 | date = May 1939 | pmid = 19873125 | pmc = 2142006 | doi = 10.1085/jgp.22.5.649 | author-link1 = Kenneth Stewart Cole }}</ref> 1907年，Louis Lapicque 提出，动作电位因膜电位跨过阈值而产生，<ref name=":17" group="lower-alpha">{{cite journal | vauthors = Lapicque L | year = 1907 | title = Recherches quantitatives sur l'excitationelectrique des nerfs traitee comme une polarisation | journal = J. Physiol. Pathol. Gen | volume = 9| pages = 620–635 }}</ref> 这后被证明是离子电导的动力系统的产物。1949年，Alan Hodgkin 和 Bernard Katz 完善了 Bernstein 的假说，他们认为轴突膜对不同的离子可能有不同的通透性；特别是，他们证明了钠通透性在动作电位中关键作用。<ref name="hodgkin_1949" group="lower-alpha">{{cite journal | vauthors = Hodgkin AL, Katz B | title = The effect of sodium ions on the electrical activity of giant axon of the squid | journal = The Journal of Physiology | volume = 108 | issue = 1 | pages = 37–77 | date = March 1949 | pmid = 18128147 | pmc = 1392331 | doi = 10.1113/jphysiol.1949.sp004310 | author-link1 = Alan Lloyd Hodgkin | author-link2 = Bernard Katz }}</ref> 他们首次实际记录了神经元膜上的介导动作电位的电学变化。<ref name=":0" group="lower-Greek">{{cite journal |last=Warlow|first=Charles| name-list-style = vanc |title=The Recent Evolution of a Symbiotic Ion Channel in the Legume Family Altered Ion Conductance and Improved Functionality in Calcium Signaling|journal=Practical Neurology|volume=7|issue=3|pages=192–197|url=http://pn.bmj.com/content/7/3/192.full|publisher=BMJ Publishing Group|access-date=23 March 2013|url-status=live|archive-url=https://web.archive.org/web/20120314104408/http://pn.bmj.com/content/7/3/192.full|archive-date=14 March 2012|df=dmy-all|date=June 2007}}</ref> 这一系列的研究最终产生了 Hodgkin、Katz 和 Andrew Huxley 在 1952 年 5 篇论文，他们应用电压钳技术来确定轴突膜的钠离子和钾离子的通透性对电压和时间的依赖关系，从而能够定量地重现动作电位。<ref name="hodgkin_1952" group="lower-alpha" /> Hodgkin 和 Huxley 在数学模型中，刻画了可能存在于包括“开放”、“封闭”和“失活”的几种不同的状态的不同离子通道的性质。他们的假说在 20 世纪 70 年代中期和 80 年代得到 Erwin Neher 和 Bert Sakmann 的证实，他们发明了膜片钳（patch clamping）技术来检测单个离子通道的电导状态。<ref name="patch_clamp" group="lower-alpha">{{cite journal | vauthors = Neher E, Sakmann B | title = Single-channel currents recorded from membrane of denervated frog muscle fibres | journal = Nature | volume = 260 | issue = 5554 | pages = 799–802 | date = April 1976 | pmid = 1083489 | doi = 10.1038/260799a0 | author-link1 = Erwin Neher | bibcode = 1976Natur.260..799N | s2cid = 4204985 }}<br />* {{cite journal | vauthors = Hamill OP, Marty A, Neher E, Sakmann B, Sigworth FJ | title = Improved patch-clamp techniques for high-resolution current recording from cells and cell-free membrane patches | journal = Pflügers Archiv | volume = 391 | issue = 2 | pages = 85–100 | date = August 1981 | pmid = 6270629 | doi = 10.1007/BF00656997 | s2cid = 12014433 }}<br />* {{cite journal | vauthors = Neher E, Sakmann B | title = The patch clamp technique | journal = Scientific American | volume = 266 | issue = 3 | pages = 44–51 | date = March 1992 | pmid = 1374932 | doi = 10.1038/scientificamerican0392-44 | author-link1 = Erwin Neher | bibcode = 1992SciAm.266c..44N }}</ref> 在 21 世纪，通过原子分辨率的晶体结构，<ref name="doyle_1998" group="lower-alpha">{{cite journal | vauthors = Doyle DA, Morais Cabral J, Pfuetzner RA, Kuo A, Gulbis JM, Cohen SL, Chait BT, MacKinnon R | display-authors = 6 | title = The structure of the potassium channel: molecular basis of K+ conduction and selectivity | journal = Science | volume = 280 | issue = 5360 | pages = 69–77 | date = April 1998 | pmid = 9525859 | doi = 10.1126/science.280.5360.69 | bibcode = 1998Sci...280...69D }}<br />* {{cite journal | vauthors = Zhou Y, Morais-Cabral JH, Kaufman A, MacKinnon R | title = Chemistry of ion coordination and hydration revealed by a K+ channel-Fab complex at 2.0 A resolution | journal = Nature | volume = 414 | issue = 6859 | pages = 43–8 | date = November 2001 | pmid = 11689936 | doi = 10.1038/35102009 | bibcode = 2001Natur.414...43Z | s2cid = 205022645 }}<br />* {{cite journal | vauthors = Jiang Y, Lee A, Chen J, Ruta V, Cadene M, Chait BT, MacKinnon R | title = X-ray structure of a voltage-dependent K+ channel | journal = Nature | volume = 423 | issue = 6935 | pages = 33–41 | date = May 2003 | pmid = 12721618 | doi = 10.1038/nature01580 | bibcode = 2003Natur.423...33J | s2cid = 4347957 }}</ref> 荧光距离测量<ref name="FRET" group="lower-alpha">{{cite journal | vauthors = Cha A, Snyder GE, Selvin PR, Bezanilla F | title = Atomic scale movement of the voltage-sensing region in a potassium channel measured via spectroscopy | journal = Nature | volume = 402 | issue = 6763 | pages = 809–13 | date = December 1999 | pmid = 10617201 | doi = 10.1038/45552 | bibcode = 1999Natur.402..809C | s2cid = 4353978 }}<br />* {{cite journal | vauthors = Glauner KS, Mannuzzu LM, Gandhi CS, Isacoff EY | title = Spectroscopic mapping of voltage sensor movement in the Shaker potassium channel | journal = Nature | volume = 402 | issue = 6763 | pages = 813–7 | date = December 1999 | pmid = 10617202 | doi = 10.1038/45561 | bibcode = 1999Natur.402..813G | s2cid = 4417476 }}<br />* {{cite journal | vauthors = Bezanilla F | title = The voltage sensor in voltage-dependent ion channels | journal = Physiological Reviews | volume = 80 | issue = 2 | pages = 555–92 | date = April 2000 | pmid = 10747201 | doi = 10.1152/physrev.2000.80.2.555 }}</ref> 和冷冻电子显微的研究，<ref name="cryoEM" group="lower-alpha">{{cite journal | vauthors = Catterall WA | title = A 3D view of sodium channels | journal = Nature | volume = 409 | issue = 6823 | pages = 988–9, 991 | date = February 2001 | pmid = 11234048 | doi = 10.1038/35059188 | bibcode = 2001Natur.409..988C | s2cid = 4371677 | doi-access = free }}<br />* {{cite journal | vauthors = Sato C, Ueno Y, Asai K, Takahashi K, Sato M, Engel A, Fujiyoshi Y | title = The voltage-sensitive sodium channel is a bell-shaped molecule with several cavities | journal = Nature | volume = 409 | issue = 6823 | pages = 1047–51 | date = February 2001 | pmid = 11234014 | doi = 10.1038/35059098 | bibcode = 2001Natur.409.1047S | s2cid = 4430165 }}</ref> 研究人员开始了解这些电导状态的结构基础，以及通道对离子种类的选择性。<ref name="yellen_2002" group="lower-alpha">{{cite journal | vauthors = Yellen G | title = The voltage-gated potassium channels and their relatives | journal = Nature | volume = 419 | issue = 6902 | pages = 35–42 | date = September 2002 | pmid = 12214225 | doi = 10.1038/nature00978 | bibcode = 2002Natur.419...35Y | s2cid = 4420877 }}</ref>  

 

20 世纪是电生理的重要时期。Julius Bernstein 于 1902 和 1912 两次提出动作电位是由轴突膜对离子的渗透性的改变引起的假说。<ref name="bernstein_1902_1912" group="lower-alpha" /> Ken Cole 和 Howard Curtis 证实了 Bernstein 的假说，他们发现在动作电位期间膜电导增加。<ref name=":16" group="lower-alpha" /> 1907年，Louis Lapicque 提出，动作电位因膜电位跨过阈值而产生，<ref name=":17" group="lower-alpha" /> 后来被证明为离子电导的动力学系统的结果。1949年，Alan Hodgkin 和 Bernard Katz 完善了 Bernstein 的假说，他们认为轴突膜对不同的离子可能有不同的通透性；特别是，他们证明了钠通透性对动作电位的关键作用。<ref name="hodgkin_1949" group="lower-alpha" /> 。他们首次实际记录了神经元膜上的电变化，这些电变化介导了动作电位。<ref name=":0" group="lower-Greek" /> 这一系列的研究在 Hodgkin，Katz 和 Andrew Huxley 的 5 篇 1952 年的论文中达到了顶峰，他们应用电压钳技术来确定轴突膜对钠离子和钾离子的通透性对电压和时间的依赖性，从而能够定量地重建动作电位。<ref name="hodgkin_1952" group="lower-alpha" /> 。Hodgkin 和 Huxley 将其数学模型的性质与离散离子通道相关联，离散离子通道可以存在于几种不同的状态，包括“开放”、“封闭”和“失活”。他们的假设在20世纪70年代中期和80年代得到 Erwin Neher 和 Bert Sakmann 的证实，他们发明了膜片钳技术来检测单个离子通道的电导状态。<ref name="patch_clamp" group="lower-alpha" /> 。在21世纪，通过原子分辨率晶体结构，<ref name="doyle_1998" group="lower-alpha" /> 研究人员开始了解这些电导态的结构基础，以及离子种类的通道选择性，<ref name="yellen_2002" group="lower-alpha" /> 荧光距离测量<ref name="FRET" group="lower-alpha" /> 和冷冻电子显微研究。<ref name="cryoEM" group="lower-alpha" />  

也是 Julius Bernstein 首次将能斯特方程（[[Nernst equation]] ）引入描述跨膜静息电位；David E. Goldman 在 1943 年将这个方程推广到了以他的名字命名的戈德曼方程（[[Goldman equation]]）。<ref name="goldman_1943" group="lower-alpha" /> 钠钾泵在 1957 年被鉴定出来 <ref name=":18" group="lower-alpha">{{cite journal | vauthors = Skou JC | title = The influence of some cations on an adenosine triphosphatase from peripheral nerves | journal = Biochimica et Biophysica Acta | volume = 23 | issue = 2 | pages = 394–401 | date = February 1957 | pmid = 13412736 | doi = 10.1016/0006-3002(57)90343-8 }}</ref><ref name=":1" group="lower-Greek">{{cite press release | url = http://nobelprize.org/nobel_prizes/medicine/laureates/1997/press.html | title = The Nobel Prize in Chemistry 1997 | publisher = The Royal Swedish Academy of Science | year = 1997 | access-date = 2010-02-21 | url-status = live | archive-url = https://web.archive.org/web/20091023003257/http://nobelprize.org/nobel_prizes/medicine/laureates/1997/press.html | archive-date = 23 October 2009 | df = dmy-all }}</ref>，其性质逐渐被阐明，<ref name="hodgkin_1955" group="lower-alpha">{{cite journal | vauthors = Hodgkin AL, Keynes RD | title = Active transport of cations in giant axons from Sepia and Loligo | journal = The Journal of Physiology | volume = 128 | issue = 1 | pages = 28–60 | date = April 1955 | pmid = 14368574 | pmc = 1365754 | doi = 10.1113/jphysiol.1955.sp005290 | author-link1 = Alan Lloyd Hodgkin }}</ref><ref name="caldwell_1960" group="lower-alpha">{{cite journal | vauthors = Caldwell PC, Hodgkin AL, Keynes RD, Shaw TL | title = The effects of injecting 'energy-rich' phosphate compounds on the active transport of ions in the giant axons of Loligo | journal = The Journal of Physiology | volume = 152 | issue = 3 | pages = 561–90 | date = July 1960 | pmid = 13806926 | pmc = 1363339 | doi = 10.1113/jphysiol.1960.sp006509 }}</ref><ref name="caldwell_1957" group="lower-alpha">{{cite journal | vauthors = Caldwell PC, Keynes RD | title = The utilization of phosphate bond energy for sodium extrusion from giant axons | journal = The Journal of Physiology | volume = 137 | issue = 1 | pages = 12–3P | date = June 1957 | pmid = 13439598 | doi = 10.1113/jphysiol.1957.sp005830 | s2cid = 222188054 }}</ref> 最终用 X 射线晶体学（[[X-ray crystallography]]）测定其原子分辨率的结构。<ref name="Na_K_pump_structure" group="lower-alpha">{{cite journal | vauthors = Morth JP, Pedersen BP, Toustrup-Jensen MS, Sørensen TL, Petersen J, Andersen JP, Vilsen B, Nissen P | display-authors = 6 | title = Crystal structure of the sodium-potassium pump | journal = Nature | volume = 450 | issue = 7172 | pages = 1043–9 | date = December 2007 | pmid = 18075585 | doi = 10.1038/nature06419 | bibcode = 2007Natur.450.1043M | s2cid = 4344526 }}</ref> 相关的离子泵的晶体结构也已经被解析，为理解这些分子机器的工作原理提供了更广阔的图景。<ref name=":19" group="lower-alpha">{{cite journal | vauthors = Lee AG, East JM | title = What the structure of a calcium pump tells us about its mechanism | journal = The Biochemical Journal | volume = 356 | issue = Pt 3 | pages = 665–83 | date = June 2001 | pmid = 11389676 | pmc = 1221895 | doi = 10.1042/0264-6021:3560665 }}</ref>

也是 Julius Bernstein 首次将能斯特方程（[[Nernst equation]] ）引入描述跨膜静息电位；David E. Goldman 在 1943 年将这个方程推广到了以他的名字命名的戈德曼方程（[[Goldman equation]]）。<ref name="goldman_1943" group="lower-alpha" /> 钠钾泵在 1957 年被鉴定出来 <ref name=":18" group="lower-alpha">{{cite journal | vauthors = Skou JC | title = The influence of some cations on an adenosine triphosphatase from peripheral nerves | journal = Biochimica et Biophysica Acta | volume = 23 | issue = 2 | pages = 394–401 | date = February 1957 | pmid = 13412736 | doi = 10.1016/0006-3002(57)90343-8 }}</ref><ref name=":1" group="lower-Greek">{{cite press release | url = http://nobelprize.org/nobel_prizes/medicine/laureates/1997/press.html | title = The Nobel Prize in Chemistry 1997 | publisher = The Royal Swedish Academy of Science | year = 1997 | access-date = 2010-02-21 | url-status = live | archive-url = https://web.archive.org/web/20091023003257/http://nobelprize.org/nobel_prizes/medicine/laureates/1997/press.html | archive-date = 23 October 2009 | df = dmy-all }}</ref>，其性质逐渐被阐明，<ref name="hodgkin_1955" group="lower-alpha">{{cite journal | vauthors = Hodgkin AL, Keynes RD | title = Active transport of cations in giant axons from Sepia and Loligo | journal = The Journal of Physiology | volume = 128 | issue = 1 | pages = 28–60 | date = April 1955 | pmid = 14368574 | pmc = 1365754 | doi = 10.1113/jphysiol.1955.sp005290 | author-link1 = Alan Lloyd Hodgkin }}</ref><ref name="caldwell_1960" group="lower-alpha">{{cite journal | vauthors = Caldwell PC, Hodgkin AL, Keynes RD, Shaw TL | title = The effects of injecting 'energy-rich' phosphate compounds on the active transport of ions in the giant axons of Loligo | journal = The Journal of Physiology | volume = 152 | issue = 3 | pages = 561–90 | date = July 1960 | pmid = 13806926 | pmc = 1363339 | doi = 10.1113/jphysiol.1960.sp006509 }}</ref><ref name="caldwell_1957" group="lower-alpha">{{cite journal | vauthors = Caldwell PC, Keynes RD | title = The utilization of phosphate bond energy for sodium extrusion from giant axons | journal = The Journal of Physiology | volume = 137 | issue = 1 | pages = 12–3P | date = June 1957 | pmid = 13439598 | doi = 10.1113/jphysiol.1957.sp005830 | s2cid = 222188054 }}</ref> 最终用 X 射线晶体学（[[X-ray crystallography]]）测定其原子分辨率的结构。<ref name="Na_K_pump_structure" group="lower-alpha">{{cite journal | vauthors = Morth JP, Pedersen BP, Toustrup-Jensen MS, Sørensen TL, Petersen J, Andersen JP, Vilsen B, Nissen P | display-authors = 6 | title = Crystal structure of the sodium-potassium pump | journal = Nature | volume = 450 | issue = 7172 | pages = 1043–9 | date = December 2007 | pmid = 18075585 | doi = 10.1038/nature06419 | bibcode = 2007Natur.450.1043M | s2cid = 4344526 }}</ref> 相关的离子泵的晶体结构也已经被解析，为理解这些分子机器的工作原理提供了更广阔的图景。<ref name=":19" group="lower-alpha">{{cite journal | vauthors = Lee AG, East JM | title = What the structure of a calcium pump tells us about its mechanism | journal = The Biochemical Journal | volume = 356 | issue = Pt 3 | pages = 665–83 | date = June 2001 | pmid = 11389676 | pmc = 1221895 | doi = 10.1042/0264-6021:3560665 }}</ref>