《生理學(xué)》BIOELECTRICALPHENOMENONANDEXCITABILITY

1、Section II Bioelectrical Phenomenon and Excitability,馮鑒強(qiáng) 教授,Electrical potentials exist across the membranes of all cells of the body. “Excitable cells or tissues” for instant: nerve and muscle cells can generate rapidly changing electrochemical impulses at the membranes. Other cells: glandular cell

2、s, macrophage, ciliated cell etc. Local changes in membrane potentials play significant roles in controlling many of cells functions.,Electrofish,Application of bioelectrical phenomenon in clinic: EEG(腦電圖)，ECG(心電圖)， EMG（肌電圖）etc.,Measurement of cellular bioelectricity,The method for measuring the mem

3、brane potential is simple in the theory but often difficult in practice due to the small sizes of most of the fiber. Figure 2-9 shows:,When the “indifferent electrode” and recording electrode( a glass microelectrode containing an electrolyte solution ) are put on the surface of cell membranes, no po

4、tential difference is recorded between two recording sites( the potential is zero). When the recording electrode passes through the cell membrane, the potential decreases rapidly to -90 millivolts,Resting membrane potential,At the resting condition ( the cells or the nerve fibers are not stimulated)

5、, the potential difference existing between the outside and inside of the membranes is called resting membrane potential. Extent of the resting membrane potential -10100mV Means the potential inside the cell or the fiber is 10 to 100 millivolts more negative than the potential in the extracellular f

6、luid on the outside of the cell or the fiber.,The resting membrane potentials of nerve fiber and muscle cell of mammalian are about -7090mV.,Several physiological terms,Polarization(極化)：the state of resting membrane potential that is, negative potential exists inside of the membrane, positive potent

7、ial outside of the membrane Depolarization(去極化或除極化)：negative potential value inside the membrane decreases (-70mV -60mV),Hyperpolarization(超極化)：negative potential value inside the membrane increases( -70mV -90mV) Repolarization(復(fù)極化)：After depolarization, the potential recovers in the resting membran

8、e potential direction.,Action potential(AP,動(dòng)作電位),When the cell or the nerve fiber is stimulated by certain intensity of electrical stimuli, the membrane potential happens a sudden change from the normal resting negative membrane potential to a positive potential and then ends with an almost equally

9、rapid change back to the negative potential, that is the action potential.,The potential rises from -70-90mV to +20+40mV then lowers to -70-90mV. The successive stages of the action potential:,Resting stage The membrane to be “polarized”(-70mV) Depolarization stage The normal “polarized” state of -7

10、0mV rising rapidly in positive direction.,Overshoot(超射)：In large nerve fibers, the membrane potential beyond the zero level and becomes somewhat positive, but in smaller fibers and many central nervous system neurons, the potential merely approaches the zero level, does not overshoot.,Repolarization

11、 stage,After overshoot, the membrane potential returns back to the normal negative potential After potential (后電位) 1)depolarizing afterpotential (去極化后電位) or negative after potential 2)hyperpolarization after potential(超極化后電位) or positive potential,Characteristics of AP in single cell,after elicited

12、by threshold stimuli, the superthreshold stimulus does not increase the amplitude of AP AP can transfer to surrounding site The transferring extent and distance of AP are not associated with the stimulating intensity that causes AP. Both 1) and 2) are called all or non（全或無）,Mechanism of Generating B

13、ioelectricity,the mechanism of origin of resting membrane potential 1902, Berstein uses the “membrane theory” to explain the mechanism of the membrane potential key point: 1)concentration difference of K+ ion exists between the outside and inside of cell membrane; 2)at resting state, the cell membra

14、ne only has permeability to K+ ion,Equilibrium potential of K+ (鉀平衡電位),靜息時(shí)， 細(xì)胞膜主要對(duì)K+有通透性；胞內(nèi)K+濃度高于胞外K+濃度K+從胞內(nèi)流向胞外， 使胞外正電荷增多膜外正電位膜內(nèi)負(fù)電位（靜息電位）膜內(nèi)帶負(fù)電荷的蛋白分子不能流出胞外，使胞內(nèi)負(fù)電荷增多；另外，K+外流使胞外正電荷增多，阻礙K+外流， 當(dāng)這兩種因素完全抵消使K+外流的濃度差勢(shì)能時(shí)，K+從胞內(nèi)向胞外凈移動(dòng)停止，膜外正內(nèi)負(fù)的狀態(tài)穩(wěn)定（K+平衡電位）,Nernst Equation (1889) Ek = RT/ZF * ln(K+0/K+i) Ek = -8

15、7mV 1944, Hodgkin and Katz demonstrated that Ek recorded from giant axon is -77mV near to -87mV K+-Na+ “l(fā)eak” channel: through which K+ and Na+ can leak.,Factors of affecting the level of resting membrane potential,K+ concentration outside the membrane K+the potential Membrane permeability to K+ and

16、 Na+ more permeable to K+ the potential more permeable to Na+ the potential,the activity level of Na+-K+-pump it is an electrogenic pump,Initiation of the action potential,膜受刺激而興奮膜對(duì)Na+通透性對(duì)K+的通透性（由于胞內(nèi)外Na+濃度差及靜息時(shí)胞內(nèi)負(fù)電位）Na+從胞外流入胞內(nèi)胞內(nèi)負(fù)電位迅速消失出現(xiàn)膜內(nèi)正電位膜外負(fù)電位（鋒電位） Na+ equilibrium potential 流入的Na+胞內(nèi)正電荷阻止Na+的進(jìn)一步內(nèi)

17、移Na+內(nèi)流形成的正電位足以阻止Na+順濃度差內(nèi)流Na+內(nèi)流停止（Na+平衡電位）,Molecular mechanism of initiation of AP,Voltaged-gate Na+ channel: It has two gates: one near the outside of the channel called the activation gate, another near the inside called the inactivation gate Three separate states at resting state(-90mv), activatio

18、n gate is closed.,The activated state when the potential becomes less negative than during the resting state(-90 to +35mV), cause a sudden conformational change in the activation gate to the open position. Sodium ions can pour inward, the sodium permeability of membrane increases as much as 500- to

19、5000-fold.,Inactivated state: a few 10,000ths of a second after the activation gate opens, the inactivation gate closes, sodium ions no longer can pour to the inside of membrane, the potential begins to recover back toward the resting membrane state(repolarization),The inactivation gate will not re-

20、open until the potential returns either to or nearly to the original resting membrane potential level.,Voltage gated potassium channel,has a gate Two states: A. During the resting state, the gate closes B. When the potential rises from -90mV toward zero, causes a slow conformational opening of the g

21、ate, allowing increased potassium diffusion outward.,The techniques for studying ion channels,Voltage Clamp”(電壓鉗) antagonist of sodium ion channel: TTX antagonist of potassium ion channel tetraethlammonium (四乙胺) Patch Clamp(膜片鉗) Effect: recording the ion current and conductance during single channel

22、 opens, and analyzes its other functions.,Advantages: electrode smaller, not easy to damage the cell,stimulus and Excitation (刺激與興奮),the basic concepts of stimulus and excitation stimulus: the changes in internal or external environment that cause the body to response types of stimulus: Chemical sti

23、muli, electrical stimuli(easy to control),mechanical stimuli ect.,Excitation and Excitability,Excitation: Old concept: from rest to activate or weaker activity to stronger activity New concept: can produce Action potential Excitability: ability of living tissues or cells to response to external stim

24、uli New concept: ability to produce AP,Factors of eliciting excitation by stimulus,intensity threshold intensity(threshold): the minimum intensity to cause the tissues to be excited subthreshold stimulus: suprthreshold stimulus maximal stimulus: the stimulus causing the tissues occur maximal reactio

25、n,changes frequency of stimulus-time(刺激強(qiáng)度-時(shí)間變化率) relationship between stimulating intensity and time a. threshold time(閾時(shí)間)：the shortest time needed for eliciting excitation by certain intensity.the stronger the stimulus, the shorter the stimulating time, the opposite is true.,b.Rheobase(基強(qiáng)度)：the mi

26、nimum potential for producing stimulating effect by electrical current. c.Utilization time(利用時(shí))：the shortest time for causing excitation by rheobase d.Chronaxie(時(shí)值),Inward current and outward current,Inward current causes the resting membrane potential increasesuperpolarization outward current cause

27、s the potential reducedepolarization, higher excitability.,Local excitation and Action potential,local excitation 閾下刺激膜少量Na+通道開放小量Na+內(nèi)流小的去極化+電刺激引起的去極化（疊加）膜局部去極化局部興奮（局部反應(yīng)）,Characteristics of local excitation,the stronger the stimulus, the bigger the amplitude of reaction can not transmit to longer di

28、stance in non-declined transmission manner electrotonic propagation(電緊張性擴(kuò)布) has no refractory period,can sum(總和) spatial summation(空間總和) temporal summation(時(shí)間總和) threshold potential(閾電位) （Fig.2-16） 閾電位比靜息電位負(fù)值小10-20mV 閾電位一定數(shù)量Na+通道開放Na+內(nèi)流，膜進(jìn)一步去極化更多Na+通道開放再生性循環(huán)（正反饋）Na+內(nèi)流更多Na+平衡電位形成AP的上升支,local current(

29、局部電位) Action potentials transmit on an unmyelinated nerve fiber (Fig) Action potential transmit on a myeliated nerve fiber(Fig) Salatory conduction(跳躍傳導(dǎo)) Advantage:,Changes in excitability of the cell during the excitation and recovery states,absolute refractory period Na+channel inactivation, excit

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