1. Show that you can reproduce the plots below, and be aware of unites-ohms, henrys, farads are in MKS units (volts, amps, seconds etc). In particular, when we find L in terms of and , which are...

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The problems just about some calculations and graph plotting. The equations for calculation are provided. No other source can be used. The reference material will be provided


1. Show that you can reproduce the plots below, and be aware of unites-ohms, henrys, farads are in MKS units (volts, amps, seconds etc). In particular, when we find L in terms of and , which are expressed in units of msecs, we need to convert to seconds. Hints: 2. For this question, do not include the Na+ channel. Evaluate the system at V=0 (Vm= Vr) From the equations for L, r and the equations for Q and 0, one way to have a higher 0 and Q is to increase [dn∞/dV]v. Play with the values in the and functions to produce such an effect and show that it is possible to increase Q and 0 slightly in this way. 1) Write out your new and functions, and plot out your new and , n∞ and dn∞/dV (similar to the plots from problem 1 but with your new values, the values can be any what you want, just explain or show what you did) 2) Find your L and r values at V=0 (C will uncharged, R is uncharged) 3) Calculate the new values for Q, 0 and f0 4) Show Zmag and Zphase (similar to the plots from problem 1 with your new values), and compare measured Q and 0 and f0 values to the calculated values Hints for this question: 3. Return the K+ channel parameters to their original values (from problem 1). Now include the Na+ channel, with the approximations (so that it boils down to a negative resistance, see attached materials for reference). Evaluate the system at V=0 1) Find the value for R’ [R’= (R*rm)/(R+rm)] 2) Calculate the value for Q, 0 and f0 3) Show Zmag and Zphase and compare measured Q and 0 and f0 values measured off the graph to the calculated values 4) Turtle hair cells have Q values between 2 and 5. Without changing r, L and C, calculate the value of R’ that will result in a Q value of 3. Plot Zmag and confirm the Q value measured off the graph is around 3 II. Hodgkin Huxley equations: active membrane voltage 3 membrane containing K+ channels Cell VinVout [K+]in [K+]out Vmembrane = Vm = (Vin – Vout) = Nernst potential for K+ ≡ Ek say [K+]in = 0.4M and [K+]out = 0.01M Will Vmembrane = Ek be zero, or positive, or negative? Why? also negative ions for charge balance (they are not permeable) Thought experiment: cell with membrane only permeable to K+ - in that case membrane voltage is set by [K+]. 4 membrane containing K+ channels Cell VinVout [K+]in [K+]out Vmembrane = Vm = (Vin – Vout) = Nernst potential for K+ ≡ Ek say [K+]in = 0.4M and [K+]out = 0.01M !" = $% &' () *+ , *- . !/ = -96mV Thought experiment: cell with membrane only permeable to K+ - in that case membrane voltage is set by [K+]. ~ 26mV valence: +1 for K+ also negative ions for charge balance (they are not permeable) 5 [ ] [ ]i o x X X zF RT E ln= [ ] = concentration R = ideal gas constant = 8.314 J/K°/Mole T = absolute temperature in K° (C+273~300) z = algebraic charge of the ion F = Faraday’s constant 96,500 Coulomb/mole Nernst potential ~ 26mV ENa ~ +37 mV EK ~ -90mV ECl ~ -67mV Based on these E values, is [K] greater inside or outside the cell? What about [Na], [Cl]? cell with Na+, Cl-, K+ intracellular and extracellular concentrations Cell Each ion (X) will have its own E value, based on its own concentrations and its valence. E values are not based on permeability. Vmembrane is based on E values and permeabilities. 6 Cell Vin Vout [K+]in [Cl-]in [Na+]in [Cl-]out [Na+]out [K+]out Current (I) can be applied (in hair cells, this occurs naturally via stereocilia channels, in an experiment current can be applied with an electrode) Cell membrane is lipid bilayer with capacitance ~ 1uF/cm2 I = #$ %&$ %' + )*+ &$ − -*+ + )./ &$ − -./ + )0 &1 − -1 Note: concentrations [ ] and thus E values are fixed and unchanging – they are not affected by applied current, voltage changes etc. g values do change. 7 at rest (d/dt =0), and with no injected current (I=0): 0 = #$% &' − )* + #,- &' − ),- + #. &/ − )/ resting membrane voltage ≡ &' = 012345067367508380125067508 I = :' ;&' ;< + #* &' − )* + #,- &' − ),- + #. &/ − )/ what is a simple way of thinking about the relationship between vm and the individual conductance (g) values? at rest vmembrane ~ -78mv. what does that tell us about the g values? recall: ena ~ +37 mv, ek ~ -90mv, ecl ~ -67mv explain in words how a change in g values could lead to an action potential. 8 the conductance of a given ion is modulated by having some of the channels open and others closed. the probability of voltage-gated channels being open depends on the voltage across the membrane, vm. opening occurs with a delay. therefore, the gated conductances are both voltage and time dependent. gk(v,t) gna(v,t) c m vm extracellular intracellular e na g na e k g k l g l e 9 recall: g is for “conductance” g=1/r g units are mhos (1/ohms) hodgkin & huxley model ( ) ( ) ( )kmknamnalmlmm evngevhmgevgdt dvci -×+-××+-+= 43 )1()( nn dt dn nn -+-= ab )()1( )()1( hh dt dh mm dt dm hh mm ba ba --= --= 1 1 07.0 4 1 251.0 10 30 20 18 10 25 + = ×= ×= - - = ÷ ø ö ç è æ - - - ÷ ø ö ç è æ - n n n n b a b na e e e e h h m m ( ) ÷ ø ö ç è æ - - ×= - - ×= 80 10 10 125.0 1 1001.0 n n b na e e n n these terms prescribe how the conductances for k+ and na+ vary with membrane voltage and with time: {{ gna(v,t) gk(v,t) note v is expressed in units of mv, ⍺ and β are expressed in units of ms. time & voltage dependence of k+ channel differential equation for n: αn and βn are functions of voltage: )1()( nn dt dn nn -+-= ab note, here v = (vm-vr) = actual membrane voltage - resting voltage. • probability of one gate being open = n • four gates in the k channel • all must be open for conduction • probability that a channel is open = n*n*n*n = n4 • conductance 4ngg kk ×= 11 v = (vm-vr) = membrane voltage - resting voltage. rewriting the main hh equation in terms of v: 12 talk about what would produce an offset from resting voltage. )1()( nn dt dn nn -+-= ab we will “linearize” the hodgkin-huxley model to demonstrate frequency tuning. na+ conductance is not necessary to demonstrate the basic frequency tuning. hodgkin huxley model, excluding na channel; focusing on k+ channel. α and β are functions of v 13 iii. derive electrical resonance -- voltage oscillations and tuned electrical impedance -- in squid axon by linearizing hodgkin huxley equations. (mauro et al. paper.) -65 -64 -63 -62 -61 -60 -59 0 0.5 1 1.5 2 2.5 3 time (msec) m em br an e vo lta ge (m v) ringing response in squid axon tuned impedance in squid axon same behavior as in the tuned lrc circuits from last week: 14 basic concept of “linearization”: f = f(x) (f is a function of x) taylor series around some x point, xo: 22 2 ( )( ) ( ) ( ) ......... 2 ( ) ( ) ( ) o o o o o o o o x xdf d f f x f x x x dx dx df f x f x x x dx - = + - + + » + - f xo x f(xo) slope = df/dx|o linear approximation: f (x) is ~ a straight line as long as you don’t move too far from point where you find the slope of the line. o df f x dx d d= 15 familiar linear equations: v=ir, dv/dt=i/c, v=ldi/dt voltage and current (v and i) are linearly related, through r, l, c. (there is no nonlinear term like i2, sin(i) etc.) linear equations often good approximation for “small enough” departures from starting point even if for larger departures there is a nonlinearity (consider example, y=sin(x)). -65 -64 -63 -62 -61 -60 -59 0 0.5 1 1.5 2 2.5 3 time (msec) m em br an e vo lta ge (m v) voltage “scales linearly” with current when linear approximation holds – linear approx was good for voltage departures up to 3 mv here. voltage response with current pulse of 1,2,3 and 4 na simulation of full nonlinear hodgkin huxley equations displayed linear behavior with “small enough” current pulses. when current pulses got too large, runaway nonlinear response (action potential) occurred. 16 back to hodgkin-huxley focusing on the k+ channel this is a nonlinear equation because of the voltage dependence of the probability of channel opening, n. )1()( nn dt dn nn -+-= ab to linearize, consider small perturbations ! around starting voltage, v 17 ( ) (1 ) (3)n n dn n n dt b a= - + - ( )4 34 (2)m l k v v k d vi c g v g n v n n v v dt dd d d dé ù= + + + -ë û [ ]( ) n n nn n v v v v v d d dn i n v dv dv dv a b ad w b a d é ùé ùæ ö æ ö æ ö+ + = - +ê úê úç ÷ ç ÷ ç ÷ è ø è ø è øë ûë û (1 )n nnv v nv +="" #*="" &'="" −="" )*="" +="" #,-="" &'="" −="" ),-="" +="" #.="" &/="" −="" )/="" what="" is="" a="" simple="" way="" of="" thinking="" about="" the="" relationship="" between="" vm="" and="" the="" individual="" conductance="" (g)="" values?="" at="" rest="" vmembrane="" ~="" -78mv.="" what="" does="" that="" tell="" us="" about="" the="" g="" values?="" recall:="" ena="" ~="" +37="" mv,="" ek="" ~="" -90mv,="" ecl="" ~="" -67mv="" explain="" in="" words="" how="" a="" change="" in="" g="" values="" could="" lead="" to="" an="" action="" potential.="" 8="" the="" conductance="" of="" a="" given="" ion="" is="" modulated="" by="" having="" some="" of="" the="" channels="" open="" and="" others="" closed.="" the="" probability="" of="" voltage-gated="" channels="" being="" open="" depends="" on="" the="" voltage="" across="" the="" membrane,="" vm.="" opening="" occurs="" with="" a="" delay.="" therefore,="" the="" gated="" conductances="" are="" both="" voltage="" and="" time="" dependent.="" gk(v,t)="" gna(v,t)="" c="" m="" vm="" extracellular="" intracellular="" e="" na="" g="" na="" e="" k="" g="" k="" l="" g="" l="" e="" 9="" recall:="" g="" is="" for="" “conductance”="" g="1/R" g="" units="" are="" mhos="" (1/ohms)="" hodgkin="" &="" huxley="" model="" (="" )="" (="" )="" (="" )kmknamnalmlmm="" evngevhmgevgdt="" dvci="" -×+-××+-+="43" )1()(="" nn="" dt="" dn="" nn="" -+-="ab" )()1(="" )()1(="" hh="" dt="" dh="" mm="" dt="" dm="" hh="" mm="" ba="" ba="" --="--=" 1="" 1="" 07.0="" 4="" 1="" 251.0="" 10="" 30="" 20="" 18="" 10="" 25="" +="×=" ×="-" -="÷" ø="" ö="" ç="" è="" æ="" -="" -="" -="" ÷="" ø="" ö="" ç="" è="" æ="" -="" n="" n="" n="" n="" b="" a="" b="" na="" e="" e="" e="" e="" h="" h="" m="" m="" (="" )="" ÷="" ø="" ö="" ç="" è="" æ="" -="" -="" ×="-" -="" ×="80" 10="" 10="" 125.0="" 1="" 1001.0="" n="" n="" b="" na="" e="" e="" n="" n="" these="" terms="" prescribe="" how="" the="" conductances="" for="" k+="" and="" na+="" vary="" with="" membrane="" voltage="" and="" with="" time:="" {{="" gna(v,t)="" gk(v,t)="" note="" v="" is="" expressed="" in="" units="" of="" mv,="" ⍺="" and="" β="" are="" expressed="" in="" units="" of="" ms.="" time="" &="" voltage="" dependence="" of="" k+="" channel="" differential="" equation="" for="" n:="" αn="" and="" βn="" are="" functions="" of="" voltage:="" )1()(="" nn="" dt="" dn="" nn="" -+-="ab" note,="" here="" v="(Vm-Vr)" =="" actual="" membrane="" voltage="" -="" resting="" voltage.="" •="" probability="" of="" one="" gate="" being="" open="n" •="" four="" gates="" in="" the="" k="" channel="" •="" all="" must="" be="" open="" for="" conduction="" •="" probability="" that="" a="" channel="" is="" open="n*n*n*n" =="" n4="" •="" conductance="" 4ngg="" kk="" ×="11" v="(Vm-Vr)" =="" membrane="" voltage="" -="" resting="" voltage.="" rewriting="" the="" main="" hh="" equation="" in="" terms="" of="" v:="" 12="" talk="" about="" what="" would="" produce="" an="" offset="" from="" resting="" voltage.="" )1()(="" nn="" dt="" dn="" nn="" -+-="ab" we="" will="" “linearize”="" the="" hodgkin-huxley="" model="" to="" demonstrate="" frequency="" tuning.="" na+="" conductance="" is="" not="" necessary="" to="" demonstrate="" the="" basic="" frequency="" tuning.="" hodgkin="" huxley="" model,="" excluding="" na="" channel;="" focusing="" on="" k+="" channel.="" α="" and="" β="" are="" functions="" of="" v="" 13="" iii.="" derive="" electrical="" resonance="" --="" voltage="" oscillations="" and="" tuned="" electrical="" impedance="" --="" in="" squid="" axon="" by="" linearizing="" hodgkin="" huxley="" equations.="" (mauro="" et="" al.="" paper.)="" -65="" -64="" -63="" -62="" -61="" -60="" -59="" 0="" 0.5="" 1="" 1.5="" 2="" 2.5="" 3="" time="" (msec)="" m="" em="" br="" an="" e="" vo="" lta="" ge="" (m="" v)="" ringing="" response="" in="" squid="" axon="" tuned="" impedance="" in="" squid="" axon="" same="" behavior="" as="" in="" the="" tuned="" lrc="" circuits="" from="" last="" week:="" 14="" basic="" concept="" of="" “linearization”:="" f="f(x)" (f="" is="" a="" function="" of="" x)="" taylor="" series="" around="" some="" x="" point,="" xo:="" 22="" 2="" (="" )(="" )="" (="" )="" (="" )="" .........="" 2="" (="" )="" (="" )="" (="" )="" o="" o="" o="" o="" o="" o="" o="" o="" x="" xdf="" d="" f="" f="" x="" f="" x="" x="" x="" dx="" dx="" df="" f="" x="" f="" x="" x="" x="" dx="" -="+" -="" +="" +="" »="" +="" -="" f="" xo="" x="" f(xo)="" slope="df/dx|o" linear="" approximation:="" f="" (x)="" is="" ~="" a="" straight="" line="" as="" long="" as="" you="" don’t="" move="" too="" far="" from="" point="" where="" you="" find="" the="" slope="" of="" the="" line.="" o="" df="" f="" x="" dx="" d="" d="15" familiar="" linear="" equations:="" v="IR," dv/dt="I/C," v="LdI/dt" voltage="" and="" current="" (v="" and="" i)="" are="" linearly="" related,="" through="" r,="" l,="" c.="" (there="" is="" no="" nonlinear="" term="" like="" i2,="" sin(i)="" etc.)="" linear="" equations="" often="" good="" approximation="" for="" “small="" enough”="" departures="" from="" starting="" point="" even="" if="" for="" larger="" departures="" there="" is="" a="" nonlinearity="" (consider="" example,="" y="sin(x))." -65="" -64="" -63="" -62="" -61="" -60="" -59="" 0="" 0.5="" 1="" 1.5="" 2="" 2.5="" 3="" time="" (msec)="" m="" em="" br="" an="" e="" vo="" lta="" ge="" (m="" v)="" voltage="" “scales="" linearly”="" with="" current="" when="" linear="" approximation="" holds="" –="" linear="" approx="" was="" good="" for="" voltage="" departures="" up="" to="" 3="" mv="" here.="" voltage="" response="" with="" current="" pulse="" of="" 1,2,3="" and="" 4="" na="" simulation="" of="" full="" nonlinear="" hodgkin="" huxley="" equations="" displayed="" linear="" behavior="" with="" “small="" enough”="" current="" pulses.="" when="" current="" pulses="" got="" too="" large,="" runaway="" nonlinear="" response="" (action="" potential)="" occurred.="" 16="" back="" to="" hodgkin-huxley="" focusing="" on="" the="" k+="" channel="" this="" is="" a="" nonlinear="" equation="" because="" of="" the="" voltage="" dependence="" of="" the="" probability="" of="" channel="" opening,="" n.="" )1()(="" nn="" dt="" dn="" nn="" -+-="ab" to="" linearize,="" consider="" small="" perturbations="" !="" around="" starting="" voltage,="" v="" 17="" (="" )="" (1="" )="" (3)n="" n="" dn="" n="" n="" dt="" b="" a="-" +="" -="" (="" )4="" 34="" (2)m="" l="" k="" v="" v="" k="" d="" vi="" c="" g="" v="" g="" n="" v="" n="" n="" v="" v="" dt="" dd="" d="" d="" dé="" ù="+" +="" +="" -ë="" û="" [="" ](="" )="" n="" n="" nn="" n="" v="" v="" v="" v="" v="" d="" d="" dn="" i="" n="" v="" dv="" dv="" dv="" a="" b="" ad="" w="" b="" a="" d="" é="" ùé="" ùæ="" ö="" æ="" ö="" æ="" ö+="" +="-" +ê="" úê="" úç="" ÷="" ç="" ÷="" ç="" ÷="" è="" ø="" è="" ø="" è="" øë="" ûë="" û="" (1="" )n="" nnv="" v="">
Answered 4 days AfterApr 03, 2022

Answer To: 1. Show that you can reproduce the plots below, and be aware of unites-ohms, henrys, farads are in...

Vijay Kumar answered on Apr 08 2022
94 Votes
Question 01:
1. Show that you can reproduce the plots below, and be aware of unites-ohms, henrys, farads are in MKS units (volts, amps, seconds etc). In particular, when we find L in terms of and , which are expressed in units of msecs, we need to convert to seconds.
Hints:
Solution 01:
Please type t
he following commands in MATLAB.
% Please note for both plots above variable V needs to be varied from -40 at step of 10 to +60 or etc.
clc
clear all
v= -40:1:10;
for i=1:1:length(v)

k=0.01;
h=(10-v(i));
l=exp((10-v(i))/10);
alp_n(i) = (k.*h)./l;
end
plot(v,alp_n)
xlabel("Voltage in mv");
ylabel("Values of alpha, beta and n_inf");
xlim([-40 10])
ylim([0 1])
hold on
for i=1:1:length(v)

k1=0.125;
beta_n(i) = k1*exp(-v(i)/80);
end
plot(v,beta_n)
hold on
for i=1:1:length(v)

k=0.01;
h=(10-v(i));
l=exp((10-v(i))/10);
alp_n(i) = (k*h)/l;
k1=0.125;
beta_n(i) = k1*exp(-v(i)/80);
n_inf(i)=alp_n(i)/(alp_n(i)+beta_n(i));
end
plot(v,n_inf);
legend('alpha_n','beta_n', 'n_inf')
% this is differential plot of .

Vs dV needs to be plotted.
clc
clear all
syms f(v)
k=0.01;
h=(10-v);
l=exp((10-v)/10);
alp_n(v) = (k*h)/l;
k1=0.125;
beta_n(v) = k1*exp(-v/80);
f(v) = alp_n(v)/(alp_n(v)+beta_n(v));
Df = diff(f,v)
v=-40:10:60;
fplot(Df, [-40 60])
legend(‘dn_inf/dV’);
This results into this
By referring to page three for the values.
%define them in MATLAB.
% now
%type the following command in MATLAB.
clc
clear all
C=10^-6;
syms f(v)
k=0.01;
h=(10-v);
l=exp((10-v)/10);
alp_n(v) = (k*h)/l;
k1=0.125;
beta_n(v) = k1*exp(-v/80);
f(v) = alp_n(v)/(alp_n(v)+beta_n(v));
Df = diff(f,v)
n0=double(f(0));
dn0=double(Df(0));
gk=30*10^3;
gl=0.3*10^3;
vk=-12*1e-3;
vl=10.6*1e-3;
r=-1/(gk*vk*4*((n0)^3)*dn0);
L=r/(double(alp_n(0))+double(beta_n(0)));
R_inv=gl+(gk*(n0)^4);
x1=@(f) (r+(2*pi.*f*L)*i)
x2=@(f) ((1-(4*pi*pi.*f.*f)+r*R_inv)+((2*pi.*f*L*R_inv)+(r*2*pi.*f*C))*i)
Z=@(f) (r+(2*pi.*f*L)*i)/(@f x2);
f = 0:10:200;
figure
subplot(2,1,1)
plot(f, abs(Z(f)))
title('Amplitude')
subplot(2,1,2)
plot(f, (180*angle(Z(f))))
title('Phase')
Question 02:
For this question, do not include the Na+ channel. Evaluate the system at V=0 (Vm= Vr)
From the equations for L, r and the equations for Q and 0, one way to have a higher 0 and Q is to increase [dn∞/dV]v. Play with the values in the and functions to produce such an effect and show that it is possible to increase Q and 0 slightly in this way.
1. Write out your new and functions, and plot out your new and , n∞ and dn∞/dV (similar to the plots from problem 1 but with your new values, the values can be any what you want, just explain or show what you did)
1. Find your L and r values at V=0 (C will uncharged, R is uncharged)
1. Calculate the new values for Q, 0 and f0
1. Show Zmag and Zphase (similar to the plots from problem 1 with your new values), and compare measured Q and 0 and f0 values to the calculated values
Hints for this question:
Solution
NA, CL, K+ all three needs to be consider in membrane...
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