regulation and measurement of intracellular calcium may 12, 2006
TRANSCRIPT
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Regulation and measurement of intracellular calcium
May 12, 2006
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The four units of the Ca signaling network. Stimuli act by generating Ca-mobilizing signals that act on various ON mechanisms to trigger an increase in the intracellular concentration of Ca. The increased level of Ca stimulates various Ca-sensitive processes to trigger many different cellular pathways. The response is terminated by OFF mechanisms that restore Ca to its resting level.
Nature reviews (MCB) 2000, 1, 11- 21
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Nat. Rev. MCB (2003) 4, 552
The regulation of intracellular calcium compartmentalization
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The four units of the Ca signaling network. Stimuli act by generating Ca-mobilizing signals that act on various ON mechanisms to trigger an increase in the intracellular concentration of Ca. The increased level of Ca stimulates various Ca-sensitive processes to trigger many different cellular pathways. The response is terminated by OFF mechanisms that restore Ca to its resting level.
Nature reviews (MCB) 2000, 1, 11- 21
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Ca ATPase2Ca/1ATP
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Ca ATPase (Ca pump)2Ca/1ATP
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Structure of the catalytic alpha subunit of the muscle Ca-ATPase
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K
Na Ca
K
H
NaGlucose
NaCaH
Ca
Glucose
H
Cl
Cl
HCO3
HCO3
lysosome
ER
mitochondria
Na = 145 mMCa = 2 mM
K = 150 mMCa = 0.1 μM
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The free energy change of moving solute X, from one side of a membrane to theotherside with concentration of C1 and C2, respectively, is Delta G = RT ln C2/C1 = 2.3 RT log C2/C1
C1 < C2, delta G is positive, need energy to occur, so called “active transport”C1 > C2, delta G is negative, spontaneously occur, “passive diffusion” or “passiveTransport”
For example, 10 fold concentration gradient across the membraneDelta G = 2.3 RT log 0.1/1 = -1,359 cal/mol (energy released in the process)
If X is a charged compound, both the chemical concentration and the electric Potential have to be consider.Delta G = 2.3 RT log C2/C1 + ZF delta V
Z: the number of chargeF: the Faraday constantDelta V: the difference of electric
potential across the membrane
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ATP
Na
K
Na
Ca
Ouabaindigitoxigenin
3Na/1CaElectrogenicReversible
- - - - - - -
+ + + + +
Na
Ca
Na = 145 mM
Na = 5 mM
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Nat. Rev. MCB (2003) 4, 552
The regulation of intracellular calcium compartmentalization
![Page 14: Regulation and measurement of intracellular calcium May 12, 2006](https://reader030.vdocuments.net/reader030/viewer/2022032703/56649d0c5503460f949e0402/html5/thumbnails/14.jpg)
The four units of the Ca signaling network. Stimuli act by generating Ca-mobilizing signals that act on various ON mechanisms to trigger an increase in the intracellular concentration of Ca. The increased level of Ca stimulates various Ca-sensitive processes to trigger many different cellular pathways. The response is terminated by OFF mechanisms that restore Ca to its resting level.
Nature reviews (MCB) 2000, 1, 11- 21
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Nat. Rev. MCB (2003) 4, 552
The regulation of intracellular calcium compartmentalization
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Ion channelsVoltage-gated ion channelsLigand-gated ion channels
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Annu. Rev. Cell Dev. Biol. (2000) 16: 521-555
Skeletal muscle
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Annu. Rev. Cell Dev. Biol. (2000) 16: 521-555
Cardiac muscle
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Membrane 去極化超過 threshold Channel 才 open
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Structure and function of the voltage-gated ion channels.
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Nat. Rev. MCB (2003) 4, 552
The regulation of intracellular calcium compartmentalization
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Acetylcholine receptor
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Structure of the acetylcholine receptor ion channel.
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An action potential is generated about every 4 ms.Action potentials move down the axonat speeds up to 100 meters per second.Their arrival at a synapse causes releaseof neurotransmitters that bind toreceptors in the postsynaptic cells,generally depolarizing the membrane(making the potential less negative) and tending to induce an action potential on it.
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The threshold potential for generation of an action potential in a postsynaptic cell.
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ATPKNa
K
Na
K
Na CaAch
Ca Secretion contraction
Memb.potential
Ligand-gated ion channelVoltage-gated ion channel
EK = - 91 mVENa = 64 mV
[K] = 140 mM[Na] = 145 mM
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Nat. Rev. MCB (2003) 4, 552
The regulation of intracellular calcium compartmentalization
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Activate PKC
Release calcium from ER
Hormone-activated phospholipase C and IP3
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IP3 receptor was cloned in 1989.
Ryanodine receptorIP3 receptor
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Molecular and Cellular Biology (1999) 190, 185-190
IP3 receptor
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Molecular and Cellular Biology (1999) 190, 185-190
Ryanodine receptor
Voltage gated Ca channel
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Sequential activation of gated ion channels at a neuromuscular junction.1 voltage-gated Ca channel2 ligand-gated nicotinic receptors3 voltage-gated Na channel to generated action potential4 voltage-gated Ca channel and Ca induced Ca release channel (ryanodine receptor)
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Role of voltage-gated and ligand-gated ion channels in neural transmission
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Annu. Rev. Cell Dev. Biol. (2000) 16: 521-555
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Troponin and tropomyosin block the interaction between myosin and actin.Troponin C: binds CaTroponin I: bind actinTroponin T: bind tropomyosinTropomyosin: double helix polypeptide
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Cardiac muscleCa entry (L channel)Ca induced Ca release from SR via ryanodine receptor
Skeletal muscle L channel directly activates Ca release from SR, extracellular Ca is not required.
Smooth musclemyosin light chain phosphorylation catalyzed by Ca/CaM MLC kinase.
+ Ca
- Ca
Myosin binding site exposed.
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neuron
myocyte
Na
Na
L (skeletal muscle)
(cardiac muscle)
L
nonexcitable cell
CaR G E
IP3
Cadepletion
(SOC)
(VOC)Ca
Na
(ROC)
Na
IP3
Ca
cAMP
Ryanodine receptor
Ca
Ca
Ca
RGE RGE
(IP3 receptor)
(IP3 receptor)
(1983)(1996)
?
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Nat. Rev. MCB (2003) 4, 552
The regulation of intracellular calcium compartmentalization
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Capacitative Ca entry
Nature reviews (MCB) 2000, 1, 11- 21
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Depletion of intracellular calcium stores activates a calcium current in mast cells. Nature (`1992) 355, 353-356.Inositol 1,3,4,5-tetrakisphosphate activates an endothelial Ca-permeable channel. Nature (1992) 355, 356-358.
Emptying of intracellular Ca2+ stores releases a novel small messenger that stimulates Ca2+ influx. Nature (1993) 364, 809-814.Depletion of InsP3 stores activates a Ca and K current by means of a phosphatase and a diffusible messenger. Nature (1993) 364, 814-818
Ca influx factor (CIF)
Activation of store-operated Ca current in xenopus oocytes requires SNAP-25 but not a diffusible messenger. Cell (1999) 98, 475-485.Store-operated Ca entry: evidence for a secretion-like coupling model. Cell (1999) 98, 487-499
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Cell (1999) 98, 487
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Cell (1999) 99, 5
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TINS (2002) 23:63-70
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Nat. Rev. MCB (2003) 4, 552
The regulation of intracellular calcium compartmentalization
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Science’s STKE 2004, January 13
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Science’s STKE 2004, January 13
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Biochim. Biophys. Acta (2004) 1742:119-131
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The four units of the Ca signaling network. Stimuli act by generating Ca-mobilizing signals that act on various ON mechanisms to trigger an increase in the intracellular concentration of Ca. The increased level of Ca stimulates various Ca-sensitive processes to trigger many different cellular pathways. The response is terminated by OFF mechanisms that restore Ca to its resting level.
Nature reviews (MCB) 2000, 1, 11- 21
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Elements of the Ca signaling toolkit.
Nature reviews (MCB) 2000, 1, 11- 21
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Catalytic domain
T286CaM-inhibitory
associationC
CaM kinase
Catalytic subunit
InhibitorycAMP cAMP
PKA
Catalytic domain
inhibitory Lipid/CaPKC
N
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Calcium-calmodulin complex mediates many cellular responses.
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N CCatalytic domain
Inhibitory domain
Calmodulin binding
Association domain
0%100%
100%
20 – 80%
20 – 80%
thr286
Thr305, 306
CaMKII
Ca/CaM
Ca increase
Ca decrease
Ca decrease
ATP
- CaM
(trapped)
(capped)
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Calcium frequency decoding mechanism by CaM-K II autophosphorylation
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Biochem. J. (2002) 364, 593-611
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Calcium regulationProteins involved in Ca signaling (mobilization)
“off” Ca pump (plasma membrane and endoplasmic reticulum)Na/Ca exchanger
“on” Ca entry (VOC, ROC and SOC)Ca release (IP3 receptor and ryanodine receptor)
Calcium sensitive cellular functionsecretioncontractionactivating calmodulin and CaMKII
Role of mitochondria in calcium regulationCalcium measurement
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[Ca]i
Ca
CaATP
IP3
Saponine or digitonin is used to permeabilized cells.
Ca electrode
IP3 was shown to be the second messenger to induce calcium release.Nature (1983) 306, 67-68
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Fura-2 acidFura-2/acetoxymethyl (AM) (membrane permeable)
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Ratioing method fura-2 + Ca ---- > fura-2-Ca (Cf) (Cb)Kd = Ca x Cf/CbCb = Cf x Ca/Kd F = SCS = I Φεl while I, incident intensity, Φ, quantum yield, ε, extinction coefficient, and l, length of light path.C = concentration of fura-2F1 (fluorescence at 340 nm) = Sf1Cf + Sb1CbF2 (fluorescence at 380 nm) = Sf2Cf + Sb2CbR = F1/F2 = (Sf1Cf + Sb1Cf x Ca/Kd) / (Sf2Cf + Sb2Cf x Ca/Kd)Ca = Kd x [(R – (Sf1/Sf2)/(Sb1/Sb2) – R] x (Sf2/Sb2)While Ca = 0, no Cb, R = Sf1/Sf2 designated RminWhile Ca = saturating conc., no Cf, R = Sb1/Sb2 designated RmaxCa = Kd x [(R – Rmin)/(Rmax – R)] x (Sf2/Sb2)Sf2/Sb2 = F of fura-2 at zero Ca (380 nm) / F of fura-2 at saturating Ca (380 nm)
= Sf2 Cf / Sb2 Cb
Ca = Kd x [(R – Rmin)/(Rmax – R)] x (Sf2/Sb2)
Peak at 340 nm, Ca under saturating conc.Ca-bound
Peak at 365nm, Ca = 0unbound
340 380360 (isosbestic point)
Excitation 340 and 380 nmEmission 505 nm
JBC (1985) 260, 3440-3450
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Calcium sensors
Ratiometric dyes
Non-ratiometric dyes
Nat. Rev. MCB (2003) 4, 579
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Protein-based calcium sensors
Nat. Rev. MCB (2003) 4, 579
aequorin
cameleon
camgaroo
pericam
coelenterazine
Tyr145 146
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Nat. Rev. MCB (2003) 4, 579
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Nat. Rev. MCB (2003) 4, 579
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Nat. Rev. MCB (2003) 4, 579