פיזיולוגיה הרצאה א - 14-02-2011
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א' ם ד א ה ל ש ה י ג ו ל ו י ז י פ ו ה י
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. . .troen@agri huji ac ilר ד ח י) ל י ד נ ק ן ( ש י ש ד" ג ן י נ 112ב
ג' ם ו י ש, א ר מ ם ו א ת ב ה: ל ב ק ת ו ע 10-12ש
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Nutrition and Neuroscience
" - ". . .Christiaan Eijkman Nobel Lecture Nobelprize org 13 Feb 2011
:// . / / / / / -http nobelprize org nobel_prizes medicine laureates 1929 eijkman.lecture html
o b e l P r i z e i n P h y s i o l o g y o r M e d i c i n e 1 9 2 9
h ris ti aa n E ij km an“ ”For his discovery of Anti Neuritic Vitamin
ir Frederick Gowland Hopkins“ -For his discovery of the growth stimulating vitam
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Nobel Prizes in NutritionScience
Nobel Prize in ChemistrySynthesis of Vitamins
Walter Norman Haworth (1937) Vitamin C
Paul Karrer (1937) Vitamin E
Robert Burns Woodward (1965)* Vitamin B 12
Structure of Vitamins
Paul Karrer (1937) Vitamin A and B
Richard Kuhn (1938) Vitamin B 2
Lord (Alexander R.) Todd (1957)*
Vitamin B 12
Dorothy Crowfoot Hodgkin (1964)*
Vitamin B 12
Nobel Prize in Physiology or MedicineDiscovery of Vitamins
Christiaan Eijkman (1929)
Vitamin B 1
Sir Frederick Gowland Hopkins (1929) Growth Stimulating Vitamins
George Hoyt Whipple (1934)*
Vitamin B 12George Richards Minot (1934)*
Vitamin B 12William Parry Murphy (1934)*
Vitamin B 12
Henrik Carl Peter Dam (1943) Vitamin K
Isolation of Vitamins
Adolf Otto Reinhold Windaus (1928)* Vitamin D
Albert von Szent-Györgyi Nagyrapolt(1937)
Vitamin C
Richard Kuhn (1938) Vitamin B 2 and B 6
Edward Adelbert Doisy (1943)
Vitamin K
" ". . .The Nobel Prize and the Discovery of Vitamins Nobelprize org 13 Feb 2011:// . / / / / / .http nobelprize org nobel_prizes medicine articles carpenter index html
( )obel Peace Prize 1949 - ( )Lord John Boyd Orr of Brechin, ; , ; )ntist MD Director of Rowett Institute Scotland Director General of UN FAO
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The Human Brain
• ~100 Million Neurons, ~ 1000
million cells• 2% Body Weight• 15% Cardiac Output• 20-30% Resting Metabolic Rate• No Stora e Tissue
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Nerve Cells
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Outline
• Membrane Potential• Resting Potential• The Nerve Cell• Action Potentials• Glial Cells• Central Nervous System
• Autonomic Nervous System – Sympathetic – Parasympathetic
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Cell Membranes and Transporters
+ ++ + +
--
-
++
+
+ +- - - -
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The cell membrane
• Diffusion• Osmosis
• Passive Transprt• Active Transport• The ionic basis of membrane
potential
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E xcita b le M e m b ra n e s
• Resting potential• Presynaptic potential – depolarization
and hyperpolarization• Ion Channels• Propagating action potentials
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Transport across cellmembranes
X+ X+
in out
• Simple diffusion• Facilitated Diffusion• Primary Active Transport• Co-transport• Counter-transport• (Endocytosis)
GRADIENT
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Simple Diffusion
GR A D I E N T
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Simple Diffusion Permeable membrane
J ( / )mmolsec = P ( / )cmsec A (cm2) (CA- CB) ( / )mmolL = ; = ; = ; =J net rate of diffusion P permeability A surface area for diffusion C concentration
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Facilitated Diffusion–Semi permeable selective membranet 0
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Facilitated Diffusion–Semi permeable selective membranet ∞
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Diffusion
Transport
Rate
Concentration
S i m p l
e d i f f u s
i o n
- Carrier mediated transport T m
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Osmosis Semi permeable membranet 0
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Osmosis Semi permeable membranet ∞
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Osmosis Semi permeable membrane
π (atm) = g (Osm/mol) C(mmol/L) σ (0-1) RTπ
= ; = / ; = ;osmotic pressure g N particles mol solution C concentration σ
=reflection coefficie
= ( . - / - ); =R gas constant 0 82 L atm mol K T absolute temperature K
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Tonicity / Osmolarity[ ] = /Urea 2 mol L[ ] = /NaCl 1 mol L
+
- -
+-
+
π (atm) = g (Osm/mol) C(mmol/L) σ (0-1) RTπ
= ; = / ; = ;osmotic pressure g N particles mol solution C concentration σ
=reflection coefficie
= ( . - / - ); =R gas constant 0 82 L atm mol K T absolute temperature K
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Tonicity / Osmolarity[ ] = /Urea 2 mol L[ ] = /NaCl 1 mol L
+
- -
+-
+
s m o l a r i t y = ( / ) g number particles mol C ( / ) concentration mol L
: / = /NaCl 2 x 1 mol L 2 Osm L: / = /Urea 1 x 2 mol L 2 Osm L
Solutions are i s o s m o t i c
π (atm) = g (Osm/mol) C(mmol/L) σ (0-1) RTπ
= ; = / ; = ;osmotic pressure g N particles mol solution C concentration σ
=reflection coefficie
= ( . - / - ); =R gas constant 0 82 L atm mol K T absolute temperature K
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Tonicity / Osmolarity[ ] = /Urea 2 mol L[ ] = /NaCl 1 mol L
To n i c i t y = π (atm) = g (Osm/mol) C(mmol/L) σ (0-1) RT
RT 37⁰C = .25 45 L • /atmmolσ NaCl : .0 3 π NaCl = 2 x 1 mol/L x 0.3 = RT = 0.6RT = 15.3 atmσ Urea : .0 05 π Urea = 1 x 2 mol/L x 0.05 = RT = 0.1RT = 2.5 atm! Solutions are o t i s o t o n i c
π
+
- -
+-
+
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Tonicity / Osmolarity[ ] = /Urea 2 mol L H y p o t o n i c[ ] = /NaCl 1 mol L H y p e r t o n i c
To n i c i t y = π (atm) = g (Osm/mol) C(mmol/L) σ (0-1) RT
RT 37⁰C = .25 45 L • /atmmolσ NaCl : .0 3 π NaCl = 2 x 1 mol/L x 0.3 = RT = 0.6RT = 15.3 atmσ Urea : .0 05 π Urea = 1 x 2 mol/L x 0.05 = RT = 0.1RT = 2.5 atm! Solutions are o t i s o t o n i c
π H20
+
- -
+-
+
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Electrogenic Transport–Semi permeable selective membrane
+
-
+- +
+
-
+- +
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Membrane Potential–Selective membrane permeable to K +, impermeable to Cl -
+- +
- +-
+
-
+-
+-
+-
+-
-V+
0
[K+Cl -] in >
[K+Cl -] out Concentration gradient
t 0
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Membrane Potential–Selective membrane permeable to K +, impermeable to Cl -
+-
+
-+ -
+-
+ -
+
-
+ -
+
-
-V+
0
[K+][Cl -] in > [K+][Cl -] out
Concentration gradient
=
( )Potential voltage gradien
t ∞
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Active Transport: Na+/K+ATPase
Na
Na
Na
Na+
K+
K+
Na+
Na+
K+
K+
K+
K+
K+
K+
K+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
K+
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Na+/K+ ATPase
Na
Na
Na
ATP
Na+
K+
K+
Na+
Na+
K+
K+
K+
K+
K+
K+
K+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
K+
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Na+/K+ ATPase
Na
ADP
P
Na+
K+
K+
Na+
Na+
K+
K+
K+
K+
K+
K+
K+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
K+
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Na+/K+ ATPase
Na
Na+
K+
K+
Na+
Na+
K+
K+
K+
K+
K+
K+
K+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
K+
Na+
Na+
ADP
P
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Na+/K+ ATPase
Na
K+
K+
K+
K+
K+
K+
K+
K+
K+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
K+
Na+
Na+
K+
K+
P
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Na+/K+ ATPase
Na
K+
K+
K+
K+
K+
K+
K+
K+
K+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
K+
Na+
Na+
P
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Na+/K+ ATPase
K+
K +
K+
K+
K+
K+
K +
K+
K+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
K+
Na+
Na+
Pi
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Na+/K+ ATPase
K++
K+
K+
K+
K+K +
K+
K+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
K+
Na+
Na+
Pi
K +
K +
K +
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Na+/K+ ATPase
K++
K+
K+
K+
K+K +
K+
K+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
K+
Na+
Na+
Pi
K +
K +
K +
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Na+/K+ ATPase
E
-E P -E P
-E PATP
+ADP Pi
3Na+in 3Na+out
2K+out2K
+in
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Transport across cellmembranes
Simple diffusion +Na+Na
+K
K+
+ + + + + + +
- - - - - - - -
- ( )Co transport Symport
+Na
GlucoseGlucose
Facilitated Diffusio
CO2
02
Cl -
Cl -
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Electrochemical Equillibrium
X+ X+
)(][][
ln)X)X)X out in
out
in
out in E E zF X
X
RT −+=(−(=(∆ +
+
+++
µ µ µ
in out
Concentration gradient
=
( )Potential voltage gradien
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Equilibrium Potential
s t Equation : E x= - . / (2 3 RT zF log 10 [ ]X in /[ ]XE = equilibrium potential ( )mV. / =2 3RT F constant (~ 60 mV at 37 ⁰ )C=Z charge on the ion (+1 for Na + & K+; - 1 for Cl -; + 2
for Ca +2 .)etc
Membrane Potential( )oldman Hodgkin Katz GHK Equat ion :
iCaiCli NaiK
oCaoClo NaoK 10m [Ca] p[Cl] p[Na] p[K] p
[Ca] p[Cl] p[Na] p[K] plog60V
+++
+++=
=P permeability to ion current
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Normal High PotassiumIon Extracellular Intracellular Extracellular Intracellular Na+ 117 30 114 29K+ 3 90 6 91Cl- 120 4 120 7.9A- 0 116 0 112.1Relative Volume 1 1.035Equillibrium potential z E xNa+ 1 35 1 36K+ 1 -89 1 -71Cl- -1 -89 -1 -71A-Membrane Potential -85 mV -68 mV
Normal Low ChlorideIon Extracellular Intracellular Extracellular Intracellular Na+ 117 30 117 30.5K+ 3 90 3 89.5Cl- 120 4 60 2A- 0 116 60 118Relative Volume 1 0.98Equillibrium potential z E xNa+ 1 35 1 35K+ 1 -89 1 -88Cl- -1 -89 -1 -89A-Membrane Potential -85 mV -85 mV
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Nerve Cells
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Nerve Cell Anatomy
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Recording Nerve Activity
, .Stuart GJ Sakmann B Active propagation of somatic action potentials into neocortical
.pyramidal cell dendrites. ; ( ): - . : .Nature 1994 Jan 6 367 6458 69 72 PMID 8107777
-V+
0
+ + ---- - + + ++ + +- - ++++- - - - -- - - - -
-85 mV
Vm
t imestimulus
h f
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Characteristics of ActionPotentials
Stereotypical size and shapePropagationAll-or-none responseVm ( )V
( )Time msec
-70
-90
0
+35
Stimulus
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Action Potentials
DepolarizationHyperpolarizationInward CurrentOutward Current
Threshold
OvershootUndershoot (hyperpolarizingafterpotential)
Refractory period
V m (mV)
-70
-90
0
+35
Stimulus
i b i f h i
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Ionic basis of the ActionPotential
1.Resting membrane potential2.Upstroke of the action potential3.Repolarization4.Hyperpolarizing afterpotential
These steps are mediated by thecollective action of specializedchannels acting together
The action potential is the summedmembrane potential of ion fluxes
enerated b all of the channels
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Gated Channels
igand Gated oltage Gated
- - -
+ + +
Rest
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Gate Channels
igand Gated oltage Gated
- - - + +
+ + +
( )Upstroke sub threshold
+Na
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Gated Channels
igand Gated oltage Gated
++++
- - - - -
( )Upstroke breakthrough threshold
+Na
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Gated Channels
igand Gated oltage Gated
++ - -
- - + +
( )Upstroke breakthrough threshold
+Na
+K
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Gated Channels
igand Gated oltage Gated
- - - -
+ ++ +
+Repolarization overshoot
+K
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Gated Channels
igand Gated oltage Gated
- - -
+ + +
Rest
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G ti f th A ti P t ti l
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potentialin
millivolts
0
-70resting state
Na + gate
opening
K+ gate
opening
Na + /K+ pump
operating
Action Potential
The action potential ispropagated down thelength of a nerve cell.
time
Generation of the Action Potential
intracellular environment of thenerve
cellmembrane
cellmembrane
[Na +] [K+]
This ion inbalance is maintained by the Na +-K+ ATPase pump.
The intracellular side of the membrane is ~70mv lower inpotential that the extracellular side.
-70mv
extracellular environment
e fr ac to ry Pe ri od
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V m (mV)
-70
-
0
+35
Equilibrium potential
Equilibrium potential
Resting Membranepotential
e fr ac to ry Pe ri od AbsoluteRelative
+Inward Na Current
+Outward K Current
.0 .0
+65
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Myelinated Axons
• Oligodendrocytes and Schwann cells• Saltatory nerve conduction• Spinal Cord Injury• Demyelinating neuropathy•
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myelinated nerve cell (Schwann cell)
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Schwanncell
unmyelinated nerve cell
myelinated nerve cell
Schwann cells in the CNS
Oligodendrocite cells in the PNS
saltatory conductancenode to node
node of Ranvier
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The Synapse
• A junction between cells allowingtransmission of information carriedby action potential
• Electrical and chemical synapses• Nerve – Nerve; Neuromuscular
synapes, etc.
• Presynaptic potential• Excitatory post synaptic potential• Inhibitory post synaptic potential
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AP
axon
synaptic vesicles
presynaptic membrane
postsynaptic membranesynapse
postsynaptic receptors direct ion channels
Action Potentials Code
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Action Potentials CodeInformation
Nerve
Receptor
Stimulus
ha si c Recep tor( )a pid ly a da pti ng
onic Receptor( )lowly adapting
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A Simple Reflex
• Knee jerk, pain – Simple stereotypical response – Does not require central processing
– Integrates external stimuli andappropriate response
• How does this give rise to more
complex phenomena?
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Sensory Input
Motor Output
Summation
Information
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