Survey
* Your assessment is very important for improving the work of artificial intelligence, which forms the content of this project
* Your assessment is very important for improving the work of artificial intelligence, which forms the content of this project
CS 621 Artificial Intelligence
Lecture 34 - 08/11/05
Guest Lecture by
Prof. Rohit Manchanda
Biological Neurons - II
08.11.05
1
The human brain
Seat of consciousness and cognition
Perhaps the most complex information processing
machine in nature
Historically, considered as a monolithic information
processing machine
08.11.05
2
Beginner’s Brain Map
Forebrain (Cerebral Cortex):
Language, maths, sensation,
movement, cognition, emotion
Midbrain: Information Routing;
involuntary controls
Cerebellum: Motor
Control
Hindbrain: Control of
breathing, heartbeat, blood
circulation
Spinal cord: Reflexes,
information highways between
body & brain
08.11.05
3
Brain : a computational machine?
Information processing: brains vs computers
- brains better at perception / cognition
- slower at numerical calculations
• Evolutionarily, brain has developed algorithms
most suitable for survival
• Algorithms unknown: the search is on
• Brain astonishing in the amount of information it
processes
– Typical computers: 109 operations/sec
11 operations/sec
–
Housefly
brain:
10
08.11.05
4
Brain facts & figures
• Basic building block of nervous system: nerve
cell (neuron)
• ~ 1012 neurons in brain
• ~ 1015 connections between them
• Connections made at “synapses”
• The speed: events on millisecond scale in
neurons, nanosecond scale in silicon chips
08.11.05
5
Neuron - “classical”
• Dendrites
– Receiving stations of neurons
– Don't generate action potentials
• Cell body
– Site at which information received is
integrated
• Axon
– Generate and relay action potential
– Terminal
• Relays information to next neuronhttp://www.educarer.com/images/brain-nerve-axon.jpg
in the pathway
08.11.05
6
Membrane Biophysics: Overview
Part 1: Resting membrane potential
08.11.05
7
Resting Membrane Potential
• Measurement of potential between ICF
and ECF
ECF
ICF
_
+
-40 to -90 mV
– Vm = Vi - Vo
– ICF and ECF at isopotential separately.
– ECF and ICF are different from each other.
08.11.05
8
Resting Membrane Potential recording
• Electrode wires can not be inserted in the cells
without damaging them (cell membrane
thickness: 7nm)
ECF
_
ICF
+
-40 to -90 mV
KCl
– Solution: Glass microelectrodes (Tip diameter: 10 nm)
08.11.05
• Glass Non conductor
• Therefore, while pulling a capillary after heating, it is filled
with KCl and tip of electrode is open and KCl is interfaced
with a wire.
9
R.m.p. - towards a theory
• Ionic concentration gradients across
biological cell membrane
Mammalian muscle (rmp = -75
mV)
ECF
ICF
Cations
Na+
145 mM
12 mM
K+
4 mM
155 mM
08.11.05
ECF
ICF
Na+
109 mM
4 mM
K+
2.2 mM
124 mM
77 mM
1.5 mM
Cations
Anions
Anions
Cl-
Frog muscle (rmp = -80 mV)
120 mM
4 mM
Cl-
10
R.m.p. - towards a theory
• Ionic concentration
gradients: squid axon
(rmp = -60 mV)
ECF
• Ionic concentration
ratios
ICF
Cations
Mammal
Frog
Cations
Na+
440 mM
50 mM
Nao/Nai
12
28
K+
10 mM
400 mM
Ki/Ko
52
56
25
33
Anions
Cl08.11.05
Anions
550 mM
40 mM
Clo/Cli
11
Ionic concentration ratios across biological cell
membranes
Species,
tissue
_Ki /
Ko
Nao/Nai
Clo/Cli
14
Squid axon
40
9
Cuttlefish axon
36
11
Crab axon
38
Frog nerve
60
3
4
Frog sartorius
muscle
56
28
33
Rat cardiac
52
12
Cat cardiac
32
25
Eel electroplaque
30
18
Nitella
0.5
0.9
1.04
Paramecium
1,065
0.02
100
Plant cells
08.11.05
12
Trans-membrane Ionic Distributions
08.11.05
13
Resting potential as a K+ equilibrium (Nernst)
potential
08.11.05
14
Resting Membrane Potential: Nernst
Eqn
Nernst Equations
RT
Vm
F
Vm
RT
F
RT
Vm
F
08.11.05
[ K ]o
EK
[ K ]i
[ Na ]o
E Na
[ Na ]i
[Cl ]i
ECl
[Cl ]o
Consider values for typical
concentration ratios
EK = -90 mV
ENa = +60 mV
r.m.p. = {-60 to –80} mV
15
Goldman-Hodgkin-Katz (GHK) eqn
Resting Membrane Potential approximat ed by the GHK eq.
RT PK [ K ]o PNa [ Na ]o
Vm
ln
F
PK [ K ]i PNa [ Na ]i
If PK PNa
RT
Vm
F
[ K ] o
EK
[ K ]i
RT
Vm
F
[ Na ]o
E Na
[
Na
]
i
If PNa PK
Taking values of R,T &F and dividing throughout by PK:
[ K ]o [ Na ]o
PNa
Vm 58 log
mV
,
PK
[ K ]i [ Na ]i
Consider = V. large, v. small, and intermediate
08.11.05
16
Equivalent Circuit Model: Resting Membrane
Out
gNa
gK
Vm
ENa
Cm
EK
In
I Na Vm E Na g Na
I K Vm EK g K
At steady state, I Na I K
Therefore, Vm
08.11.05
E Na g Na EK g K
g Na g K
17
Equivalent Circuit Model including Na pump
Out
INa(p)
gK
Cm
ENa
EK
IK(p)
In
At steady state,
I Na I K I p 0
Therefore,
Vm
08.11.05
E Na g Na EK g K I p
g Na g K
18
Membrane Biophysics: Overview
Part 2: Action potential
08.11.05
19
ACTION POTENTIAL
08.11.05
20
ACTION POTENTIAL: Ionic mechanisms
08.11.05
21
Action Potential: Na+ and K+ Conductance
g K Vm , t g K n(Vm , t )
4
g Na Vm , t g Na m(Vm , t ) h(Vm , t )
3
08.11.05
22
Action Potential Propagation
• NonS decremental,
constant
velocity
R
R
R
R
R
X=0
1
2
3
4
Convex
VTh
Concave
Time
08.11.05
23
Hippocampus: location
08.11.05
24
Hippocampal Network &
Connections
08.11.05
25
Membrane Biophysics: Overview
Part 3: Synaptic transmission &
potentials
08.11.05
26
“Canonical” neurons: Neuroscience
08.11.05
27
Synapses : Chemical & Electrical
08.11.05
28
Transmission : Chemical & Electrical
08.11.05
29
Chemical Transmission
Neurotransmitter
Receptors
08.11.05
30
Postsynaptic Electrical Effects
08.11.05
31
Synaptic Integration: The Canonical Picture
Action potential: Output signal
Axon: Output line
Action potential
08.11.05
32
The Perceptron Model
A perceptron is a computing element with input
lines having associated weights and the cell
having a threshold value. The perceptron model is
motivated by the biological neuron.
Output = y
Threshold = θ
wn
08.11.05
w1
Wn-1
Xn-1
x1
33
y
1
θ
Σwixi
Step function / Threshold
Σwixi > θ
function
y
= 1 for
Σ > θ, y >1
=0 otherwise
Features of Perceptron
• Input output behavior is discontinuous and the derivative
does not exist at Σwixi = θ
• Σwixi - θ is the net input denoted as net
• Referred to as a linear threshold element - linearity because
of x appearing with power 1
08.11.05
• y=
f(net): Relation between y and net is non-linear
34
Perceptron / ANN “neuron”
Neurophysiological basis of:
a) Input Signs
b) Input Weights
08.11.05
35
Dendrites & Synapses in Real Life !
08.11.05
36
08.11.05
37
Neuron morphologies
08.11.05
38
In the Retina …
08.11.05
39