Rao 1.

6
 Consider the input impedance of a transmission line circuit, with an
applied voltage v(t) inducing an input current i(t).
i(t)

v(t) Zin

For sinusoidal excitation, we can write:

v (t ) = V0 cos(t )
i (t ) = I 0 cos(t   )     / 2,  / 2
where  is the phase difference between voltage and current. Note that  =
0 only when the input impedance is real (purely resistive).
 The time-dependent input power is given by:

P(t ) = v(t )i (t ) = V0 I 0 cos(t ) cos(t   )

=
V0 I 0
cos( )  cos(2t   ) cos x  y = cos x cos y  sin x sin y
cos x  y = cos x cos y  sin x sin y
2
cos( x  y )  cos( x  y ) = 2 cos x cos y

The power has two (Fourier) components:

V0 I 0 V0 I 0
P (t ) = cos( )  cos(2t   )
2 2
(A) an average value: A B

V0 I 0
cos( )
2 FOURIER POWER REPRESENTATION

(B) an oscillatory component with frequency 2f:

V0 I 0
cos(2t   )
2
 We can also rewrite the time-dependent current as:

i(t ) = I 0 cos(t   ) = I 0 cos cost  I 0 sin  sin t

where we have used the trigonometry formula:

cos x  y = cos x cos y  sin x sin y

This result yields an equivalent expression for the power:

P(t ) = V0 cos(t ) I 0 cos(t   ) =

= V0 cos(t ) I 0 cos(t ) cos( )  V0 cos(t ) I 0 sin(t ) sin( )
V0 I 0
= V0 I 0 cos( ) cos (t ) 
2
sin( 2t ) sin( )
 
active ( real ) power 2 
reactive power

( 2 cos x sin x = sin 2 x ) ACTIVE/REACTIVE POWER REPRESENTATION

 ACTIVE/REACTIVE POWER REPRESENTATION

V0 I 0
P(t ) = V0 I 0 cos( ) cos (t )  2
sin( 2t ) sin( )
 
2
active ( real ) power 
reactive power

ACTIVE/REACTIVE POWER REPRESENTATION:

T
1
P(t ) =  P(t )dt =
T 0
T T
1 1 V0 I 0
=  V0 I 0 cos( )cos (t )dt  
2
sin( ) sin( 2t )dt
T 0    T 0 2
1 cos 2t
2

V0 I 0
= cos( )  0 (cos 2 x = 2 cos2 x  1)
2
This result tells us that the time-average power flow is the average of the
active power. The reactive power has zero time-average, since power is
stored and completely reflected by the reactive component of the input
impedance during the period of oscillation.
 In many engineering situations, we use the root-mean-square (r.m.s.)
values of quantities. For a given signal:

v(t ) = V0 cos(t )
the r.m.s. value is defined as:
T T
1 1
=   =  (t )dt
2 2 2
Vrms V cos ( t ) dt V cos
T 0
0 0
T 0
2
1 1
= V0 0    =
2
cos ( ) d V0
2   2
1 cos 2

2 

2 / 2
This result is valid for sinusoidal signals. Each signal shape corresponds
to a specific coefficient (peak factor = V0 /Vrms ) that allows one to convert
directly from peak to r.m.s. values.
 For a non-sinusoidal periodic signal, we can determine the r.m.s. value
by using a very important theorem of vector spaces. If we decompose the
non-sinusoidal signal into its Fourier components:
V (t ) = V0 av  V1 (t )  V2 (t )  V3 (t )   = Vk (t )
k
Then:
2

V (t )2rms =  Vk (t )  =  Vk (t )rms

2

 k  rms k

So, the r.m.s. value of the signal is computed as:

Vrms = V02av  (V1 ) 2rms  (V2 ) 2rms  (V3 ) 2rms  

 So far, we have used peak values for the amplitude of voltage and
current. In terms of r.m.s. values, the time-average power for a
sinusoidal signal is: v (t ) = V0 cos(t )
i (t ) = I 0 cos(t   )     / 2,  / 2
V0 I 0 V0 I 0
P(t ) = cos( ) = cos( ) = Vrms I rms cos( )
2 2 2
Finally, we can relate the time-average power to the phasors of
voltage and current. Since:
v(t ) = V0 cos(t ) = ReV0 exp(it )
i (t ) = I 0 cos(t   ) = ReI 0 exp(i ) exp(it )
we have phasors:

V = V0 and I = I 0 exp( i )

The time-average power in terms of phasors is given by:

P(t ) = ReV  I * = ReV0  I 0 exp(i )

1 1
2 2
1
= V0  I 0 cos( )
2
 The time-average power in terms of phasors is given by:

P(t ) = ReV  I * = ReV0  I 0 exp(i )

1 1
2 2
1
= V0  I 0 cos( )
2
Note that one must always use the complex conjugate of the phasor
current to obtain the time-average power. It is important to remember this
when voltage and current are expressed as functions of each other. Only
when the impedance is purely resistive, I = I* = I0 since  = 0.

Also, note that the time-average power is always a real positive quantity
and that it is not the phasor of the time-dependent power. It is a common
mistake to think so.
 When we define the time-average power in terms of phasor voltage and

P(t ) = ReV  I *
current as:
1
2
There may be an ambiguity if you start from the following time-dependent
quantities:
V (t ) = V0 cos(t )  phasor : V = V0
I (t ) = I 0 cos(t   )  phasor : I = I 0 exp(i )
Why should we use the complex conjugate of the current? In fact,

P(t ) = ReV  I * = ReV0  I 0 exp(i ) = V0  I 0 cos( )

1 1 1
2 2 2
but also:
P(t ) = ReV  I  = ReV0  I 0 exp(i ) = V0  I 0 cos( )
1 1 1
2 2 2
We find the same answer because:
Reexp(i ) = Reexp(i ) = Recos( )  i sin( )
= Recos( )  i sin( ) = cos( )
 Does this imply that:

ReV  I *= ReV  I 

1 ? 1

2 2
Let’s consider the more general formulation:
V (t ) = V0 cos(t  V )  phasor : V = V0 exp(iV )
I (t ) = I 0 cos(t   I )  phasor : I = I 0 exp(i I )
where  = V   I and

P(t ) = ReV  I * = ReV0 exp(iV )  I 0 exp(i I )

1 1
2 2
1 1
= V0 I 0 cos(V   I ) = V0 I 0 cos( )
2 2
but

ReV  I  = ReV0 exp(iV )  I 0 exp(i I ) =

1 1 1 1
V0 I 0 cos(V   I )  V0 I 0 cos( )
2 2 2 2
1
2
 1

Re V  I = Re V0 exp(iV )  I 0 exp(i I )
*

2
 1
2
1
= V0 I 0 cos(V   I ) = V0 I 0 cos( )
2
 ReV  I *= ReV  I 
1 ? 1

2 2
In a transmission line, we will see that both the voltage phasor and the
current phasor may be complex quantities involving a space-dependent
phase factor and other complex factors like a reflection coefficient and
characteristic impedance.

Therefore, it is essential to use the conjugate of the current phasor when

evaluating the time-average power in general. So, we use:

ReV  I *
1
P(t ) =
2
 Now, we consider power flow, including explicitly the generator, to
understand under which conditions maximum power transfer to a load
can take place.
ZG Iin
ZR
Vin = VG
Z R  ZG
1
I in = VG ZR
Z R  ZG VG ~ Vin
1

Pin = Re Vin  I in*
2

Generator Load
 RG Iin

1
2

P(t ) = Re Vin  I in*  VG ~ Vin RR
As a first case, we examine resistive impedances:

ZG = RG Z R = RR Generator Load

Voltage and current are in-phase at the input. The time-average power
dissipated by the load is:
1
2
 * 1

P(t ) = Re Vin  I in = VG
RR
2 RR  RG
*
VG
1
RR  RG
Vin = VG
ZR
Z R  ZG
1
I in = VG
Z R  ZG
1 2 RR
= VG
2 RR  RG 2
To find the load resistance that maximizes power transfer to the load for a
given generator we impose:

d P(t )  d 2 P(t ) 
= 0  and 2
 0 
dRR  dRR 
 RG Iin

1
2

P(t ) = Re Vin  I in*  VG ~ Vin RR
To find the load resistance that maximizes power
transfer to the load for a given generator we impose:
Generator Load
d P (t )
=0
dRR
from which we obtain:
1 RR
P(t ) = VG2
2 RR  RG 2
d  RR 
=0=
 
RR  RG  2 RR RR  RG 
2

 2
dRR  RR  RG   RR  RG 4
 RR  RG   2 RR = 0  RR = RG

Therefore, we conclude that for maximum power transfer the load

resistance must be identical to the generator resistance.
 ZG Iin

Let’s now consider complex impedances: i.e.

VG ~ Vin ZR

Z R = RR  iX R ZG = RG  iX G
Generator Load
We infer that for maximum power transfer, generator and load impedances
must be complex conjugates of each other since:
  
*
 Vin = VG
ZR
1
  1 
Pin = Re Vin  I in = ReVG
2
* ZR
VG 
*

2  ZG  Z R  ZG  Z R  
1
  Z R  ZG
1
I in = VG
  Z R  ZG

 
 
1 2  RR  iX R 
= VG Re 2 * 
2 (
  R  X 2
)  ( R 2
 X 2
)  Z Z *
 Z Z
R G 
G
 G
 R
 R G R
 ZG
2
ZR
2 
1 2  RR  iX R 
= VG Re 2 
2  G
( R  X 2
G )  ( RR
2
 X 2
R )  2 ( R R
G R  X R G 
X )
1 2  RR  iX R 
= VG Re 2
2  G
( R  RR ) 2
 ( X R  X G ) 
 ZG Iin

We infer that, for maximum power transfer, the VG ~ Vin ZR

generator and load impedances must be the complex
conjugates of each other since:
Generator Load
1 2  RR  iX R 
1
 
Pin = Re Vin I in = VG Re
*

 2
  2
2 2  G
( R RR ) ( X R X G ) 
1 2 RR
= VG
2 ( RG  RR ) 2  ( X R  X G ) 2
 RG = RL and X R = XG or
 Z R = Z G*
This can be easily understood by considering that, to maximize the active
power supplied to the load, voltage and current of the generator should
remain in phase. If the reactances of the generator and load are opposite
and cancel each other along the path of the current, the generator will only
see a resistance. Therefore, the voltage and current will be in-phase with
maximum power delivered to the load.
 Thus, the total time-average power supplied by the generator (under
conditions of maximum power transfer) is: ( Z R = Z G*  RR = RG and X R =  X G )

Ptot
1
2
 *

2

1 2 1
= Re VG  I in = VG
1 2 1
= VG
2 RR 4 RR
( Z R = Z G*  RR = RG and X R = XG )
and the time-average power supplied to the load is: 1
Pin = VG2
RR
2 ( RG  RR )  ( X R  X G ) 2
2

  
*
*
Pin = ReVin  I in  = ReVG
1 1  ZR 1
*
VG   
2 2  ZG  Z R  ZG  Z R  
 
1 2  RR  iX R  1 2 1 Iin
= VG Re  = VG
ZG
2
2  4 RR  8 RR
or half that! (This is VERY disappointing.) ZR
VG ~ Vin

Generator Load
 SUPPLIED BY
GENERATOR:
1
2
 *

2

1 2 1
Ptot = Re VG I in = VG
1 2 1
= VG
2 RR 4 RR
1
 
1 2 1
SUPPLIED TO
LOAD: Pin = Re Vin I in = VG
*

2 8 RR
The power dissipated by the internal generator impedance is therefore:
1
2
 *

4

1 2 1 1 2 1 1 2 1
PG = Re (VG  Vin ) I in = VG  VG
RR 8
= VG
RR 8 RR
We conclude that, in conditions of maximum power transfer, only HALF of the
total active power supplied by the generator is actually used by the load. The
generator impedance dissipates the remaining half of the available active
power. Z I
G in

This may seem a disappointing result, but it is the best

one can do for a real generator with a given internal
impedance! VG ~ Vin ZR

Generator Load
 A typical engineering problem involves the transmission of a signal from
a generator to a load. A transmission line is the part of the circuit that
provides the direct link between generator and load.

Transmission lines can be realized in a number of ways. Common

examples are the microstrip line (below), parallel-wire line and coaxial
cable. For simplicity, in most diagrams we use the parallel-wire line to
represent circuit connections, but the theory applies to all types of
transmission lines.

Generator Load

ZG
VG Transmission line ZR
~
 EXAMPLES OF TRANSMISSION LINES:
Coaxial cable
Two-wire line 2a

2a
2d

  b
Z0 = cosh1
d Z0 = ln
2a
2 a 2 a
2b
  2 
  14  8  8  1  intrinsic impedance:
 14   1
0  4h  e r 4h e r 4h  er 
Z0 = ln 1     2

2 2(1  e r )  weff  11 weff   

=
11 weff 2 
  w
    
  
t e

er
Microstrip
 If you are only familiar with low frequency circuits, you are used to
treating all lines connecting the various circuit elements as perfect wires,
with no voltage drop and no impedance associated to them (i.e. lumped
impedance circuits). This is a reasonable procedure as long as the length
of the wires is much smaller than the wavelength of the signal. At any
given time, the measured voltage and current are the same for each
ZR
location on the same wire.
VR = VG
ZG  Z R
Generator Load

ZG
VG ~ VR VR VR ZR

L << 
 For context, let’s look at some examples. The electricity supplied to
households in the USA consists of high power sinusoidal signals with
frequency of 60Hz (in the USA or 50Hz, depending on the country).
Assuming that the insulator between wires is air (e  e0), the wavelength for
60 Hz is:
c 2.999 108 m  s 1
f = c   = = 1
= 5.0  10 6
m = 5,000 km
f 60 s
which is the about the distance between San Francisco and Boston!

Let’s compare this to frequencies in the microwave range, for instance

60 GHz (used in a collision avoidance apparatus). The wavelength is given
by:
c 2.999 108 m  s 1
= = 9 1
= 5.0  10 3
m = 5.0 mm
f 60 10 s
which is comparable to the size of a microprocessor chip.
What conclusions do you draw?
 At sufficiently high frequencies, the wavelength becomes comparable
to the length of conductors in a transmission line. The signal propagates
as a wave of voltage and current along the line, because it cannot
change instantaneously at all locations. Therefore, we cannot neglect the
impedance properties of the wires (i.e. it is a distributed impedance
circuit).

V ( z) = V  eiz  V eiz
Generator Load

ZG
VG ~ V( 0 ) V( z ) V( L ) ZR

L
 Note that the (Thévenin) equivalent circuit of a generator consists of an
ideal alternating voltage generator in series with its actual internal
impedance. When the generator is open ( Z R  ) we have:

I in = 0 and Vin = VG

If the generator is connected to a load Z R: Iin

ZR ZG
Vin = VG
ZG  Z R
VG
1
I in = VG ~ Vin ZR
ZG  Z R
If the load is a short ( Z R = 0 ):
VG
Vin = 0 and I in = Generator Load
ZG
 The simplest circuit problem that we can study consists of a voltage
generator connected to a load through a uniform transmission line. In
general, the impedance seen by the generator is not the same as the
impedance of the load, because of the presence of the transmission line,
except for some very particular (peculiar) cases.

Z in = Z R
Z in Transmission line ZR
only if

L=n (n integer)
2
L
Our goal is to determine the equivalent impedance seen by the generator,
that is, the input impedance of the line terminated by the load. Once that is
known, standard circuit theory can be used.
Generator Load
ZG

VG Transmission line ZR

Generator Equivalent Load

ZG

VG Zin
 A uniform transmission line is a “distributed circuit” that we can
describe as a cascade of identical cells with infinitesimal length. The
conductors used to realize the line possess a certain series inductance and
resistance. In addition, there is a shunt capacitance between the
conductors, and even a shunt conductance if the medium insulating the
wires is imperfect. (We use the concept of shunt conductance, rather than
resistance, because it is more convenient for adding the parallel elements
of the shunt.) We can represent the uniform transmission line with the
distributed circuit below (so-called general lossy line): i.e.

L dz R dz L dz R dz

C dz G dz C dz G dz

dz dz
 The impedance parameters L, R, C, and G represent:
L = series inductance per unit length;
R = series resistance per unit length;
C = shunt capacitance per unit length;
G = shunt conductance per unit length

Each “cell” of the distributed circuit will have impedance elements with
values: Ldz, Rdz, Cdz and Gdz, where dz is the infinitesimal length of the
cells.

If we can determine the differential behavior of an elementary cell of the

distributed circuit, in terms of the voltage and current, we can then find a
global differential equation that describes the entire transmission line. We
can do so because we assume the line to be uniform along its length.

So, all we need to do is to study how voltage and current vary in a single
elementary cell of the distributed circuit.
L dz R dz L R
dz dz

C dz G dz C dz G dz

dz dz
 In many cases, it is possible to neglect resistive effects in the line. (In
this approximation there is no Joule effect loss because only reactive
elements are present.) The equivalent circuit for the elementary cell of a
loss-less transmission line is shown in the figure below:

L dz
I (z) I (z)+dI

V (z) C dz V (z)+dV

dz
 The series inductance determines the variation of the voltage from input
to output of the cell, according to the sub-circuit below:
L dz

V (z) I (z) V (z)+dV

dz
di(t )
The corresponding (phaser) circuit equation is: v(t ) = L
dt
(V  dV )  V = iL  dzI
which gives a first order differential equation for the voltage: i.e.
dV
= iL  I
dz
 The current flowing through the shunt capacitance determines the
variation of the current from input to output of the cell.

I (z) I (z)+dI
dI
C dz
V (z)+dV
dv(t )
i (t ) = C
The (phaser) circuit equation for the sub-circuit above is: dt
0
dI = iC  dz(V  dV ) = iC Vdz  iC  dV  dz
The second term (including dV∙dz) tends to zero very rapidly for
infinitesimal length dz (it is the product of TWO infinitesimals) and can be
ignored, giving a first order differential equation for the current:

dI
= iC  V
dz
 Thus, we have obtained a system of two coupled, first order
differential equations that describe the behavior of voltage and current on
the uniform loss-less transmission line. The equations must be solved
simultaneously.
 dV
 dz = iL  I
 dI
 = iC  V
 dz
These are often called “telegraphers’ equations” of the loss-less
transmission line.
I (z) L dz I (z)+dI

V (z) C dz V(z)+dV

dz
 One can easily obtain a set of uncoupled equations by differentiating with
respect to the space coordinate. The first order differential terms are
eliminated by using the corresponding telegraphers’ equation: i.e.

dV d 2V dI
= iL  I  2 = iL  = iL( iC  V ) =  2 LC  V
dz dz dz
dI d 2I dV
= iC  V  2 = iC  = iC ( iL  I ) =  2 LC  I
dz dz dz

These are often called “telephonists’ equations”.

 d 2V d 2I
2
=  2
LC  V and 2
=  2
LC  I
dz dz
Now, we have two uncoupled, second order differential equations for
voltage and current, which give an equivalent description of the loss-less
transmission line. Mathematically, these are wave equations and can be
solved independently.

The general solution for the voltage equation is:

d 2V
2
=  2
LC  V  V ( z ) = V  e iz  V  e iz
dz
where the wave propagation constant is:

   LC
Note that the complex exponential terms including  have unitary
magnitude and purely “imaginary” argument, therefore they only affect the
“phase” of the wave in space.
    LC
We have the following useful relations:

2 2f   e r r
= = = = =  e r r e 0 0 =  e
 vp vp c
Here,  = v p / f is the wavelength of the dielectric medium surrounding the
conductors of the transmission line and
1 1
vp = =
e re 0 r 0 e
is the phase velocity of an electromagnetic wave in the dielectric. As you can
see, the propagation constant  can be written in many different, equivalent
ways.
 The current distribution on transmission line can be readily obtained by
differentiation of the result we obtained for the voltage: i.e.
dV  =  LC
= iL  I and V ( z ) = V  e iz  V  e iz
dz
dV
 = iV  e iz  iV  e iz = iL  I
dz
which gives:

I ( z) =
L

C  iz   iz
V e V e 
Z0

1   i z

V e  V  e  i z 
The real quantity:
L
Z0 =
C
is called the “characteristic impedance” of the loss-less transmission line.
 The solution for a uniform lossy transmission line can be obtained
following a very similar procedure, using the equivalent circuit for the
elementary cell shown in the figure below.

L dz R dz
I (z) I (z)+dI

V (z) C dz G dz V (z)+dV

dz
 The series impedance determines the variation of the voltage from input
to output of the cell, according to the sub-circuit:
L dz R dz

V (z) I (z) V (z)+dV

dz
di(t )
The corresponding circuit equation is: v(t ) = L  R  i (t )
dt
(V  dV )  V = (iLdz  Rdz)  I
from which we obtain a first order differential equation for the voltage:
dV
= (iL  R )  I
dz
 Likewise, the current flowing through the shunt admittance determines
the input-output variation of the current, according to the sub-circuit:

I (z) dI I (z)+dI

C dz G dz V (z)+dV

dv(t )
i (t ) = C  G  v(t )
The corresponding circuit equation is: dt
dI = (iC  dz  Gdz )(V  dV ) 0

= (iC  G ) Vdz  (iC  G )  dV  dz

The second term (including dV∙dz) can be ignored, giving a first order
differential equation for the current:
dI
= (iC  G )  V
dz
 We have (again) a system of coupled, first order differential
equations that describe the behavior of voltage and current on the lossy
transmission line:
 dV
 dz = (iL  R )  I
 dI
 = (iC  G )  V
 dz
These are the “telegraphers’ equations” for the case of the lossy
transmission line.
L dz R dz
I (z) I (z)+dI

V (z) C dz G dz V (z)+dV

dz
 One can easily obtain a set of uncoupled equations by differentiating with
respect to the coordinate z (as done earlier):

dV d 2V dI
= (iL  R )  I  2 = (iL  R )  = (iL  R )  (iC  G )  V
dz dz dz
dI d 2I dV
= (iC  G )  V  2 = (iC  G )  = (iC  G )  (iL  R )  I
dz dz dz

These are the “telephonists’ equations” for the lossy transmission line.
 The telephonists’ equations for the lossy transmission line are
uncoupled, second order differential equations and are (again) wave
equations. The general solution for the voltage equation is:
d 2V  z  z
2
= ( iL  R )  ( iC  G )  V  V ( z ) = V e  V e
dz
V ( z ) = V  e z  V  e z = V  e z e iz  V  e z e iz
where the wave propagation constant is now the complex quantity:

 = (iL  R )  (iC  G )    i
The real part, , of the propagation constant, , describes the attenuation of
the signal due to resistive losses. The imaginary part, , describes the
propagation properties of the signal waves as in loss-less lines.

The exponential terms including  are “real”. Therefore, they only affect the
“magnitude” of the voltage phasor. The exponential terms including  have
unitary magnitude and purely “imaginary” argument, affecting only the
“phase” of the waves in space.
 The current distribution on a lossy transmission line can be readily
obtained by differentiation of the result for the voltage:

dV
= (iL  R)  I with V ( z ) = V  e z  V  e z
dz  = (iL  R)  (iC  G )
dV
 = V  e z  V  e z = (iL  R)  I
dz
which gives:
(iC  G )
I ( z) =  (V  e z  V  e z )
(iL  R )
1
= (V  e z  V  e z )
Z0
with the “characteristic impedance” of the lossy transmission line:

(iL  R ) Note: the characteristic

Z0 = impedance is now complex!
(iC  G )
 For both loss-less and lossy transmission lines:
the characteristic impedance does not depend on the line length
but only on the metal of the conductors, the dielectric material surrounding
the conductors and the geometry of the line cross-section, which determine
L, R, C, and G.

One must be careful not to interpret the characteristic impedance as some

lumped impedance that can replace the transmission line in an equivalent
circuit.
This is a very common mistake!

Z0 ZR Z0 ZR
Find the ratio of the inner to outer conductor diameters for a coaxial cable
with a characteristic impedance of Z0 = 50 W if e = 2.56e0. (polystyrene—
no known organism degrades it and filled with air)

Likewise for a coaxial cable with a characteristic impedance of

Z0 = 75 W if e = 2.25 e0. (polyethylene, PE, or Teflon, PTFE)

Standard coaxial line impedance for RF power transmission in the U.S. is

exclusively almost 50 W.

Why?
 A neuron is the basic anatomical unit in the nervous system. There are many
extensions of the single neuronal cell that are long and filamentary; these structures are
called dendritic processes. Every neuron plays several roles in the neural system:
1. The metabolic machinery within the cell provides a
power source for information-processing functions;
2. A tree of processes called dendrites is covered with
special structures called synapses, where junctions are
formed with other neurons. These synaptic contacts are
the primary information-processing elements in neural
systems.
3. The “processes” act like wires, conveying information
over a finite distance. The resistance of the fine
dendrites allows the potential at their tips to be
computed with only partial coupling to the computations
in the tree.
4. Temporal integration of signals occurs over the short
term through charge stored on the capacitance of the
cell membrane, and over the longer term by means of
internal secondary messengers and other complex A conceptualization of a classic neuron. Synpatic inputs are collected by
biochemistry. the dendritic tree and are integrated on the capacitance of the cell body.
5. Some neurons are equipped with a long specialized When the potential at the axon hillock exceeds threshold, a nerve pulse
is generated and is propagated down the axon. The capacitance of the
process called an axon that is used for digitizing data for axon to the extracelluar fluid is redced by the myelin sheath. Breaks in
local transmission and for transmitting data over long the sheath (nodes of Ranvier) allow periodic restoration of the pulses..
distances.

Your brain (3 lb. weight) may contain up to one trillion neurons.

Na+ ions:
q
q
1 q 1 1 q2
V=
4e ri
 W= 
4e 0 ri
dq =
8e ri
e H 0 = 80  e 0
2

e lipid = 2  e 0
Na  r = 0.1 nm W = 100 meV ( water ) ~ 4k BT
(r = 0.117 nm W = 2.4 eV (lipid ) ~ 100k BT
membrane is a near-perfect insulator
dehydrated Na) to ion flow
 Erwin Neher and Bert Sakmann's
PATCH-CLAMP TECHNIQUE:

Vp
Glass Pipette

Rp

Rs
Vm Extracellular

Fluid
Cm Rc Rm
Upward steps due to formation of sodium
Cell Membrane Ion channels; downward steps due to the channels
Channel closing. The height of a single step is the current
Vc
Intracellular in a single channel, which increases
Fluid approximately linearly with applied voltage
 The charge-transfer in all nerve membranes is driven metabolically by pumps that
actively expel Na+ from the cytoplasm and import K+ from the extracellular fluid.
nerve membrane circuit
I = (VK  V )GK  (VNa  V )GNa  (VCl  V )GCl
Typical concentrations of ions inside neural
processes and in extracellular fluid (Katz, 1966).

• batteries represent the reversal potentials for particular ions;

• conductances represent the membrane permeability.
• membrane capacitance is shown as a lumped element

k BT N in N in for I = 0 : •• chlorine
G ~20G
= e Vr / k BT
K Na

Vr =  ln where neglected
q N ex N ex (VK  V0 )GK  (VNa  V0 )GNa = 0
( Nernst Equation ) VK GK  VNa GNa
The diffusion of ions outward will be exactly V0 = ~ 85mV
balanced by the drift inward when the voltage GK  GNa
across the membrane reaches the value Vr such
that (just like the built-in potential in a pn-diode).
resting potential
• An action potential is
triggered by the de-polarization of the
plasma membrane—that is to say, a
shift in the membrane potential to a
less negative value inside.
• De-polarization opens voltage-gated
Na+ channels, allowing a small number
of Na+ ions to enter the cell down its
electrochemical gradient.
• The influx of positive charge de-
polarizes the membrane further,
thereby opening more Na+ ion
channels, which admit more Na+ ions
causing more de-polarization.
• This self-amplification process (an
example of positive feedback)
continues until the electrical potential in
the local region of the membrane has
shifted from its resting value of about
85 mV to almost as far as the Na+
equilibrium potential of about +40 mV.
• At this point, the cell would come to a
new resting state, with all of its Na+
channels permanently open.
 Two mechanisms act in concert to
cause the sodium channels to close
again:
1. the Na+ channels have an
automatic inactivating mechanism
that causes the channels to re-
close rapidly even though the
membrane is still de-polarized.
The Na+ channels remain in this
inactivated state, unable to re-
open until the membrane
potential has returned to its initial
negative value.
2. Voltage-gated K+ channels
provide a second mechanism in
most nerve cells to help restore
the original negative potential.
These channels open in response
to membrane de-polarization, but
with slightly slower kinetics.
dVm
I = Cm  (VK  Vm )GK n 4  (VNa  Vm )GNa m3h  (VCl  Vm )GCl  (Vl  Vm )Gl
dt
 The ability to control the current into or out of one electrical node by the potential on another node
is the key ingredient that makes all information processing possible. (Transistors serve this function in
a digital computer.) The function of the synapse is to control the conductance of the membrane separating
the interior of the post-synaptic cell from the extracellular fluid.
When the pre-synaptic
membrane is de-polarized, (A) (B)
calcium channels allow
calcium ions Ca2+ to flow into
the pre-synaptic cell from the
synaptic cleft. Ca2+ activate
the sub-cellular machinery
that causes the release of
neurotransmitter molecules
into the synaptic cleft, where
these molecules diffuse to
the post-synaptic membrane
and initiate a chain of events
that results in the opening of
ion-specific channels.
 The nervous system is very different from a digital microprocessor. Generally, a
very large number of inputs are brought together or aggregated in an analog
fashion. The classic neuron is equipped with a tree of filamentary dendrites that
aggregate synaptic inputs from other neurons. The input currents are integrated by
the capacitance of the cell until a critical threshold potential is reached, at which
point an output is generated in the form of a nerve pulse—the action potential. This
output pulse propagates down the axon, which ends in a tree of synaptic contacts to
the dendrites of other neurons.
Thus, we can think of the simplest neuron
as forming the analog sum of the inputs from
the axons and dendrites of other neurons in a
tree of passive dendritic processes, such as
that shown below. The fan-in of an average
neuron ranges from 1,000-3,000, which
contrasts with the fan-in for a circuit in a digital
computer of 2-4.

A dendritic tree of a neuron. This structure may be used to generate

action-potential outputs on an axon. Synapses on the tips of the tree
provide outputs from, as well as input to the dendritic processes.
http://www.youtube.com/watch?feature=player_detailpage&v=yy994HpFudc
https://www.youtube.com/watch?v=2TIK9oXc5Wo

Simulation of neural oscillations at 10

Hz. Upper panel shows spiking of
individual neurons (with each dot
representing an individual action
potential within the population of
neurons), and the lower panel the
local field potential reflecting
their summed activity.
 We can model a dendritic process by adding capacitors at every node (just
like a transmission line.)
R dz
I (z) I (z)+dI

V (z) C dz G dz V (z)+dV

dz
The differential equations for the current and voltage are:

 dV
 = R  I
dz
 dI
 = (iC  G )  V
 dz
 One can easily obtain a set of uncoupled equations by
differentiating with respect to the coordinate z (as done earlier):

dV
= R  I d 2V dI V
dz  2 =  R  = R  (iC  G )  V = RG V  RC
dI
= (iC  G ) V dz dz t
dz
or
d 2V V
= RG V  RC generally (absent phasors)
dz 2
t V (r , t )
=   D(V , r )V (r , t )
t
This equation is called the diffusion equation; it governs the time-
course of signal propagation in a dissipative passive medium
where the stuff out of which the signal is made is stored in the
medium and the fraction lost as the signal propagates is
proportional to the amount present. Passive dendrites of a neuron
obey this equation.
 V (r , t )
=   D(V , r )V (r , t )
t

d 2V V
DIFFUSION EQUATION:
= RG  V  RC
dz 2 t
Passive dendrites of a neuron obey this equation.
• V is the voltage across the membrane;
• C is the capacitance of the membrane;
• R is the axial resistance of the cytoplasm; and
• G is the conductance to the extracellular fluid (all per unit length).

(The flow of heat in a medium is governed by the same equation with V

playing the role of temperature, C is the heat capacity and R is the
thermal resistance.)

(The diffusion of minority carriers in a semiconductor follow the same

equation as well—in this case G models recombination with majority
carriers.)
 d 2V V
DIFFUSION EQUATION: = RG  V  RC
dz 2 t
We use the method of separation of variables to solve for (transient) V:
V ( z , t ) = Z ( z )T (t )
d 2V V d 2Z T
= RG  V  RC  T = RG  ZT  RCZ
dz 2 t dz 2 t
T d 2Z 1 1 T
= RG  ZT  RCZ
ZT dz 2
ZT ZT t
 d 2Z
 =  2
Z
2
1d Z 1 T  dz 2
= = RG  RC    2
 
Z dz 2 T t T 1 2
 =   D T
 t  
where  =C /G and D = 1 /( RC )

time constant diffusionconstant
We use the method of separation of variables to solve for V:
 d 2Z where =
C/
G and D = 1 /( RC )
 2
=  2
Z  
dz time constant diffusionconstant
 T
 1 
=   D2 T V = Z ( z)  T (t ) = eiz e st
 t  
1 
T =e st
 s =   D2 
 
  = 0  T = e t / 

Z = eiz   1
s = 0   =  D   = i
2

where 
==
RG 1 /
D
space constant

 is the characteristic length for diffusion

 d 2Z
 =  2
Z where 
=
C/
G and D = 1 /( RC )
dz 2 
 T time constant diffusionconstant
 1 
=   D2 T
 t   V = Z ( z)  T (t ) = eiz e st

When we solve for V (for the case where G  0), assuming a harmonic
excitation, we find the classic form for a wave traveling at velocity :

1 2
G 0
T =e st
 s =   D   s = i =  D2

R dz
 I (z) I (z)+dI

i 1 
2 =    =  (1  i)
V (z) C dz G dz V (z)+dV

D 2 D
dz

 V = eiz eit = e z ei (t  x / ) where  = and  = 2 D
2D