6. THE ELECTRICAL SYSTEM OF THE BODYAll body functions are controlled by the electrical system! information transfer in the central and autonomous nervous system, operation of the brain and spinal cord, operation of the muscle functions operation of the body organs to control and maintain system billions of electrical signals have to be generated in the human nervous system the source of the electrical signals are electrochemical potentials in the nerve cells = measurement of electrical signals and electrical potential in nerve transmission (As well as body response) allows to obtain useful clinical information. Electromyogram (EMG) muscle function Electrocardiogram (BCG) heart function Electroencephalogram (EEG) brain functions Electroretinogram (ERG) Electrooculogram (EOG) eye functions The nervous system of the body is structured in two parts: the central nervous system controls the voluntary body functions the autonomous nervous system controls the involuntary body functions ‘The central nervous system consists of brain, spinal cord and the peripheral nerves Neurons transfer information (obtained by the acoustical, optical, temperature, odor, etc sensitive sensor system of the body) to the brain and the spinal cord. ‘They also transmit response information from the brain to the appropriate muscles. The autonomous nervous system concentrates on controlling and maintaining the functions of all the inner organs, heart and intestines. It cannot be controlled voluntarily. The infromation transfer is also controlled by neurons. 225The basis structural unit of the nervous system is the neuron It allows the transmission and reception of electrical pulses which are the basis for the information transfer. basis structure of neurons is similar in all living creatures most of the available information about neuron structure has been obtained by studying the neurons of squids The main difference is the number of neurons and the complexity of the neuron interconnections which forms a net of potential information links for example a lobster has only a few thousand neurons, the brain of a human operates with ~ 100 billion (10'!) neurons. Each of the neurons is linked to an average of 10° other neurons which gives a total of 10! connections. The figure gives an impression about the connections between the neurons. The contact points between the neurons are called the synapses.‘The single neuron is a fairly complex nerve fiber. The figure shows the structure of a motor neuron, which connects directly to a muscle. Dendnites Muscle fibres The basic parts of a neuron cell are axons, nucleus, dentrites, and synapses nucleus is the core of the cell body which contains the DNA genetic information to control the operation of the cell dentrites are receiver antennas of neurons synapses are contact points to other neurons axon is a long fiber which allows to transmit electrical signals to the final receiver. — Node of Ranvier Synapses on a neuron. 227The neuron cell operates unidirectional, the receiving end is at the dentrites either through direct contact to the sensors or through synaptic connections with other neurons. The transmitting axis is the axon which is connected to other neurons or to the receptor. The dentrites have a threshold for incoming signals, only electrical signals with an amplitude above the threshold are accepted and the signal is transmitted further along the axon. The axon in human neurons has a diameter of © 10-20 yim and can have a length of © Im. in the human it extends from the brain to low in the spinal cord or from the spinal cord to the finger ete The electrical pulse along the axon travels with a speed of 0.6 - 100 m/s. The conductivity depends mainly on the membrane of the axon be- cause the neuron operates mainly on the basis of an electrical potential difference between the interior and the exterior side of the membrane. This potential difference is caused by different concentration of negative and positive charged ions on either side. Inside of oxon Exiroceliulor fluid cole, [o*] «15 (= ] [ot ]= 145 97 kt] =150 It ]«s 03 [Misc] = 5 [er J=o fer J=125 139 Misc ]=156 [Misc] = 30 2 ve-70mVv ve0 The achieved potential difference is between AV = 60 - 90 mV. This is the resting potential of the axon, when no stimulating pulse has been received. 228Potential difference is achieved due to selective diffusion of positive ions through the membrane which causes an enrichment of positive charges outside the cell. This causes the potential difference AV ~-70 mV and therefore an electrical field E. ‘The equilibrium will be established for a certain concentration ratio which is described by the Nernst equation: CG xs ~2e(Vi-Vo)/kT a ® With the electrical charge e=1.6-10-19C, and the Boltzmann constant k= 1.38-10-*J/K, and z as the valence of the ion 13.7 for positive ions 0.073 for negative ions Membrane Pe e+ + F Extocellvar tlud Longitudinal. section Cross-section of nerve fire of nerve fibre (polr'sed) The thickness of the cell membrane is z 5-6 nm, if it is covered with myelin it is 2 pm. -70- 10-3 BY = 10 V 21.17. 10°V/m ‘The accumulated charge @ at the surface S of the membrane is: (for a dielectrical constant « £7, the electrical permittivity € = 8.85:10- C?/N - m2) Q=ke:S+E 229‘The intracellular and extracellular liquid are two conductive fluids, the membrane is an insulator => axon membrane is a cylindrical capacitor The capacitance for a membrane of thickness b is: Q_ QkeS _ Key S Cape Q:6 6b ‘The capacitance per surface area is: Key F =— # 0.01-, 5 © OTe AIQ sI0 and the charge density o at the surface of the membrane is: Cv 46 on ea eT For mylinated fibers the thickness of the membrane is much larger which reduces the capacity /area by a factor of 300.EXAMPLE Calculate the capacity of an unmyelinated axon with a lenght of L = 1m, a membran thickness of b = 10 nm, and a radius of a= 2.5 pm, Calculate the total amount of charge on the axon surface! The capacity of the axon is: Ke-S _ w-€ oat iE 8 (an-a-L) Cc =97-10° x C=9.17-10-*§ = 0.1 pF For a voltage of V = 70 mV across the membrane of the axon the total charge is: Q = C-V = 68nC For a myelinated axon the length of the myelinated section is D = 1.4 mm and its thickness is b = 2 um. This yields for the capacity C and the total charge of a myelinated sector: C = 68-10% F Q = 48-10% C This yields a total capacity and charge for a 1 m long axon of: C=49-10°F = 49nF Q = 34-10" C = 3470 capacity and charge are much smaller than for the unmyelinated axon.leakage current accross cell membrane ‘The membrane is not a perfect insulator therefore a leakage current accross the membrane will occur. The leakage current is determined by the resistance Ry, of the membrane material. hay . po With pm 1.6107 Om as electrical conduetivity of the membran material The leakage current Im discharges the capacitance, the time is determined by the time constant 7=R-C. This yields a differential equation for the potential difference v: wviesev dt Rac with the solution VQ =e" The time constant T=Rm - Cm = KEoPm 210-3 232The resistance and the capacitance of a portion of the axon membrane are expressed in terms of the axon radius r and length L using S = 2naL. KeodraL Cee _ Pmb Fm = 2naL The capacitance increases with length, the resistance decreases with length. EXAMPLE Calculate the membrane resistance R,, for the unmyelinated and myelinated axon of the previous example. For the unmyelinated 1 m long axon the resistance is: Pmb 1.6- 107 [Qm]_10-8 [m] pa a ee a yt Qra-L ~ Qn 25-10 fm] Tf) ~ 1°? Rn For the 1.4 mm long myelinated axon section the resistance is: 1.6 - 107 [Am] 2- 10-8 Ry [Om] a 2m 25- 10-8 [m) 1. This indcates a much higher resistance than for the unmyelinated axon. The time constant 7 is not affected because it is independent of the axon dimensions. T = K-€0'pm = 10-107 5resistance along the axon Because the external and internal fluids are conductive, a current can flow inside or outside of the axon. The interior fluid has a certain resistance, which is determined by the conductivity p; of the fluid and by the axon radius @ and length L. ma? what pi is the conductance of the axoplasma: pi 0.5 2m The internal resistance increases with decreasing radius a. The resistance per unit length is: R_ pi NST 3a 234When a section of the membrane is excited by electrical current or external stimulus, the membrane permeability changes, ion exchange takes place across the membrane causing a rapid change in potential according to the Nernst equation: —re(Vi-Ve)/kT Cc. This changes the polarity because positively charged ions enter the interior of the axon. The polarisation time scale is determined by the time constant r = Rm C. Because of the potential difference along the outer surface of the axon, electrical currents are induced in the internal conductive fluids. Therefore the depolarization moves along the axon. External current flow nae aN“ ere polonsed §——Yy Yepcteiseat polarised Axon Se mee WT) TTS SS Fee? =o HE ~~ ~~ ~~ oe ie i I Membr Depotarisation —_Repotarisation epaow Direction of transmission < + Action potential ipotentcl aide. wiki : respect to outside) 235‘The process can be described by application of Kirchhoff’s law for the current conditions in the axon. © current across the membran: Im =42 = Cm» % av By (Ohms law, AV is the potential change along the interior of the axon due to depolarization) © current along the inside of the axon: J; = 1dv hale Applying Kirchhoff’s Law that at a node point the sum of the incoming currents is equal to the sum of the outgoing currents: K(@) = He 82) = In = AT ~ In = Cn dividing the entire equation by the surface area element AS = 2nadz yields the cable equation Gm WV _ Im, 1 av AS dt AS * Iran; dz? 236which describes the propagation of a pulse along a ‘conductive’ material The change of the potential with time is a function of the leakage current through the surface of the membrane and the change of voltage along the axis. Capacitance of membrane be NL ford AK/\RLA)V\) TAWA) A | sh Resistance of axon The speed of the pulse is determined by the capacitance C’ and by the internal resistance R;. As larger the capacitance, as longer it takes the membrane to discharge (time constant), therefore a smaller propagation speed. However as larger the internal resistance, as smaller the axial current qj The resistance is inverse to the axon radius, as larger the radius, as higher the propagation velocity. 237EXAMPLE Longitudinal current as a function of position 1 Imm —>} 0.5mm be * The voltage along the axis is shown at some instant time. The axon radius is a=10 jum, the resistivity of the axoplasma is pi=0.5Qm. The longitudinal current J; can then be expressed as a function of position +20 — av/ax = 110%10-8/1%10-5 = 110 V m-! av/AK = -(110«10-3)/(0.54 0-5) = - 220 V m-! -90 os —= 1 a Pa ‘fa > 1dV WO)= Rae Left hand side: dV/de = 0.11 V/10-$m Right hand side: dV/dz = -0.11 V/0.5-10-'m = -220 V/m resistance per unit length r r(a) = 2 = i = 1.6 + 10°Q/m Left hand side current: I; = -7-10- A Right hand side current: J; = 14-10-§ A 238The cable equation can be reformulated: wv__m, AS 1 dv dt ~ Gm” Ir-a riCm da? with & = "8,rj = 2 and AV = Vyeak — yest & 70 mV: dv AV bea av dt oT 2pi + K+ €y dx? multiplying with the time constant rT = K- copm yields: WV _ ayy be _ ay +h88 7 R pi K- ey dx? introducing the space constant \ = 4 and using the time ik +€9* fm Yields the final formulation for the motion of the signal along the axon with time, constant T = ey av AV = Voeat = Vet = M5 = 7 for an unmyelinated axon with a=2.5 pm and with b=6 nm and a myelinated axon with b=2 yum, respectively the space constants Aym and Am are calculated to: = 49-104m Am = 89-103 m with a time constant of r = 10-3 s 239The remaining question is: What is the speed of the pulse moving along the axon? The speed of the pulse vin an axon fiber can be determined from the ratio of space constant and time constant, The speed increases proportionally with the square-root of the radius. This are severe limitations since the radius of unmyelinated human axons are typically smaller than lym. EXAMPLE fora <1ym + v $03m/s This indicates slow response times of Tresp(L=1 m) > 3.28 Even for larger sized unmyelinated axons the speeds remain slow and the response time large. fora =2.5ym > v =0.5m/s this indicates a relatively slow response, Tresp(L=1 m) = 2s. for a = 1 mm (squid) + v= 10 m/s fast response time, Tyesp(L=1 m) = 0.1 s. 240For a myelinated axon fiber there is a fixed relation between the radius of the axon a and the thickness of the myelin layer 6: 6 ~& 0.4a. This simplifies the expression for the space constant ); ya feb em - log. gz. bm 2 Bi Bi To? v= fo2-P-a = Jor POH” 4 ~ 2530-0 pi 05 this yields for the speed of the signal in an myelinated axon: vy = d= 2 06.108. a m/s T K+ €o* Pm é w% 8 E . Myelinated a B4F & 2 Unmyelinated 0 9 oO 100 200 300 400 600x10" am Myelinated 0 2 4 6 8 10x10" Inner radius, a, m The speeds increases linearly with the axon radius a.EXAMPLE For a = 0.015 mm = 15 ym > v=3.9-10' m/s 26 ys. fast response time, Tresp(L=1 m) ‘The response time is significantly faster than in the case of unmyelinated axons. Even for thin axon fibers the speed is significant. For a = 0.2 pm + v=5.1-107! m/s this results in a response time, Treap(L=1 m) = 2s. Properties of unmyelinated and myelinated axons of the same radius. Quantity Unmyelinated Myelinated ‘Axon inner radius, @ 5x10" m 5x10 m Membrane thickness 6x10"? m Myelin thickness 2x10-8 m Ke 6.20x10"" S10" mt 6.20 x 10°" sm" ‘Axoplasm resistivity 110m 119m Membrane (resting) or 107m 107 2m myelin resistivity pm Time constant 7=K¢Pm e2xtos 62x10" s Space constant d = iin ee Vi na afSben = i Bo, V~ Bp, 165Va =370x10°§ m Node spacing D Conduction speed trom amped model ~S70va Conduction speed, Upenyoinated™1800VE Unyesnates™17% 108 empirical Ratio of empirical to model 67 7.2.06 38 conduction speed ‘Space constant using thick membrane model nea (OBA 2p, fini ¥Pm nay = 12408 Pi 2426. THE ELECTRICAL SYSTEM OF THE BODY 6.2. Electrical Signals as Indicator for Body FunctionsElectrical potentials and signals are originated by pulse transmission in the neuronal sensor and motor nerve system. detection of the signals allows to test the operation of the bnerve system and the associated muscle functions! EMG electromyogram measures the electrical activity of the muscles and the associated sensor and motor nerve system! helps to check muscle operation, muscle weaknesses! ECG electrocardiogram measures electrical signals and electrical potentials associated with the operation of the heart muscle! helps to check heart conditions! EEG electroencephalogram measures the electrical activity of the brain! helps to test brain functions! ‘The electrical activity is triggered by electrical pulses moving along the nerve system as described in previous chapter. The pulse cam be either directly detected by planting a pick-up electrode near the sensor or motor nerves (see e.g. EMG). This method is not advisable for sensitive muscle and neuronal systems like heart and/or brain. Indirect methods are therefore based on checking potential differences at a ‘safe’ distance from the muscle system (ECG, EEG). 245Potential pulse is triggered by current J; along the axon and travels with speed v = X/r and pulse height AV = Vpeax — Vrest & 110 mV. The pulse duration is a few milliseconds depending on the time constant r. The potential at a distance r from axon can be expressed in terms of the current and the conductivity oo (<1/p) of the surrounding medium (co 0.08-0.33 [A/Vm]). The current is expressed in terms of the axon conductivityo; and radius a: I _ 2 a VO) = Gg with T= ana? r y ae ‘ds fa NY The current can be estimated in the framework of a simple triangular pulse model (see previous example). v ~~ 0 246‘The potential outside the axon represents an overlap from the various contributions of the pulse. P th (ath) th x 1 TD; _ (2-224) Amr - ao 1 To 12, This yields on the basis of the triangular pulse approximation: Ve= ave 2 ( Ler Iai t/a uss) Lar _ Vert Var , Ifa 4 a0 m1 ro re Typically, o/oy © 10. Using the previous example, 2; *1 mm, x2 ©0.5 mm: This results (for a¥2.5um) to potentials of approximately : - 10-3 ‘ - 19-12 2) = MM? 1825-107 Fl yy (L342) 4 Ty TO 72. 247v= 172-10-9(2-3 mT) At distances in the order of [ pen potenti ection ‘ons asiThe choice of probes define shape and pulse height of the signal! A needle electrode is closer but has a small solid angle, it therefore is preferable for monitoring pulses along only a few axon fibers. A plate electrode is planted on the surface of the skin. It warrants a larger solid angle but the detected signal reflects the total pulse integrated over many axon fibers. Pulse shape is also slightly different because of different solid angle. mv + my o ° a 1 mv a1 mv pele plate 252The electrodes can pick up voluntary signals of muscle action and response. The voluntary signals are however spread in time (100 ms) because of time differences in axon signals. To measure muscle or sensor response conditions externally stimulated signals are typically used. Soesher rave | | aT _ aes act Acton poten! Tmo occa The stimulator pulse (Vs ~ 100 V, Ats ~ 0.1- 0.5 ms) triggers a response signal in the muscle (or sensor) system which can be detected after some latency time which is due to the response time of the nerve system. Assuming a typical speed of v © 100 m/s for the signal, the delay time + between the stimulating pulse and the signal measured at a distance of 6x = 0.5 mis: Ar v 0.5 m ——— w 5-103 = 100 m/s 5-10 s 5 ms The amplification displays pulse height of “1 mV. 253By choosing various levels of stimuli pulses, different parts of the sensor and motor nerve system can be stimulated. Weak stimuli pulses are detected by the sensitive sensor nerves, the detected response pulses show a large latency because of the associated travel time of sensor pulse to brain or spine and the response motor pulse to electrode. Seine cord z over neve 3a 5 é obtrtitit 10 03 a oh a 4 S meee o eli © 10 20 30 a Tne (nic) _ Atgens + Atmor 0.5m + 1m _ a T = Trens + Tot = : © m/s 110s = 15ms Stronger stimuli pulses triger signals both in the motor and sensor nerves and both response signals will be detected at different delay times. Strong stimuli pulses trigger only the motor nerves, no detectable response from the sensor nerves. 254‘These examples show that an average pulse speed explains the observed latencies in the measured signals. A reliable test of muscle functions and response requires a better defined measurement of the associated pulse speeds. ‘The pulse speed along the motor nerve axons can be determined using two stimuli signals applied at different locations and measuring the time difference between the two associated response signals at a single electrode. Recording Stimulus Stimulus electrode 2 1 Stimulus [8 msec: The delay time between the two signals is 7 = 4 ms. Ag Ar r= ee v T 255To measure the speed of the pulse along the sensory nerve system a single stimulus signal is used and the response signal is detected with two or three electrodes. The time difference between the different response signals 71,2, 72,3, 71,3 gives the speed information. [0.15 m— [0.25 m6 20 me Sv Vi i Swmulys 1 2 Recording electroves ay ~1 mv th 27-4 sec = 7.0 mice 4 11 mee ——4 — Time Ar 0.25 m v 0.2m = Mes Tacaoss = $8m/s ms = Tapa, = 50m/s the results indicates position dependent signal velocities, 256ELECTROCARDIOGRAM ECG The electrocardiogram represents the measurement of the electrical signals associated with the pulsation of the heart muscle. Because it is not advisable and difficult to place an electrode into the heart muscle, external potential activity has to be measured at certain locations of the torso. From mans . 2 tangs snoatiat YY Se room fy ber \ avioverneuly eRe Right ventricle YY Lett ventricle The operation of the heart has been discussed in detail in section 3.2. The heart functions as a double pump system for the blood flow through the body (systemic cycle) and the lungs (pulmonary cycle). The pumping through each of these cycles is induced by contraction of the atrium muscle, which pumps the blood into the ventricles, subsequent contraction of the ventricle muscle pumpes the blood into the blood vessels. The contraction is induced by (involuntary) electrical signals, the sinoatrial node (SA) for the atrium and the atrioventricular node (AV) which induces the ventricular chambers to contract. The rate of these stimuli signals is about 72 1/min (pulse). 297The associated current flow along the heart muscle induces electrical field lines and equipotential lines (points of identical potential) perpendicular to the field lines on the surface of the torso. CG) om CD, a) current Flow and electric Field. lines en ©) Equipetentials oa the +4 ey Surface. b) Equpelential lines ona plane The equipotential lines change with time due to periodicity of the stimuli pulses. Tos chest chest back. back. 258The electrocardiogram is a representation of the surface potential at a certain position at the torso as function of time. It is measured as potential difference between three points at the chest (or back) of the torso. Ec c ex Lead II bees Uh positions are typically right shoulder (A), left shoulder (B) and left leg (C). The potentials between (A-B) V4 — Vp, (A-C)Va — Vo, and (B-C) Vp — Vo are called lead I, lead II, and lead III, respectively. Other combinations including additional resistors between the positions allow the measurement of augmented lead configurations (for details see book). 259‘Typical ECG shows potential signal from the different leads as a function of time. Signal structure indicates the operation of the sinoatrial and atrioventricular nodes. Example shows potential at lead IT as a function of time. Peak P is associated with the sinoatrial node and the subsequent contraction of the atrium chamber, peak QRS to T are associated with the atrioventricular node. Potential Time (sec) Figure shows ECG for three standard leads, the three augmented leads and six additional positions along the chest (near the heart). These last leads give directional information of the P dipole vector. i) ET reget me ae Negative slectroge Tinateent| fE wo ze 260EXAMPLES FOR HEART MISFUNCTIONS Right ventricular hypertrophy corresponds to an enlargement and thickening of the right ventricle. Because of the larger volume of the right ventricle the dipole vector P changes direction and the peak QRS in lead I turns negative. The leads V, and V2 show strong signal because the correspond to the right ventricle. Left ventricular hypertrophy cooresponds to an enlargement and thickening of the left ventricle. Mi V2 Ys % Vs Ye Na NR NA NA The thicker left ventricular wall causes the QRS dipole to point to the left, subsequently lead I turns large. Also the lead Vp turns negative and the left hand side leads Vi, V5, and Vg show strongly enhanced signals. 261A fault in the electrical conduction system of the heart is called a bundle block, it blocks and slows down the electrical signals because the have to travel through the lesser conductive surrounding muscle material. The example shows a characteristic ECG for a right bundle branch block which slows the soignal down at the right ventricle: vi ve The indication in lead Vj shows a more pronounced positive signal and in lead Vp a more pronounced and prolonged bipolar signal.ELECTROENCEPHALOGRAM EEG The electroencephalogram represents the measurement of the electrical signals associated with the function or misfunction of the brain. Again the electrodes are attached externaly as it is not advisable to plant them directly into the brain material. Positions of the electrodes depend on the goal of the EEG. Standard locations are shown in figure: Measured are potential differences between the electrodes and a reference electrode usually attached to the ear. The locations are symmetrical to test for left/right asymmetries. The typical amplitudes of brain signalsare in the range of © 30 - 50 V. The EEG is therefore handicapped by low level electrical noise. A EEG of a (supposedly normal) person is shown. ek : CyB, ee, Tyler eM lan A, meinen Re Fram Be wrayer apneThe frequency of the EEG signals depends on the mental activity of the person. A relaxed state corresponds to frequencies in the range from 3 to 13 Hz, an excited person shows frequencies above 13 Hz. The figure shows EEGs for a sleeping person. The early phase of sleep is characterized by higher frequencies than the deep-sleep phase. /-—H1 sec 100 LV} AA ret neattt Brain misfunctions can be monitored. For example severe epilepsy can be recorded by its pronounced spiky signals with large amplitudes in excess of 100 |muV. Less severe epilepsy shows more rounded voltage peaks. ho see fe) O,-Ay Grond mol () Ty-Ay 3" ony + Petit mal 264