Intracellular Ca 2+ Signaling 2004. Why Calcium? Na + and Cl - are sea water –Excluded to maintain low osmotic pressure –[K + ] i kept high for electrical.

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Presentation transcript:

Intracellular Ca 2+ Signaling 2004

Why Calcium? Na + and Cl - are sea water –Excluded to maintain low osmotic pressure –[K + ] i kept high for electrical neutrality [Ca 2+ ] i maintained very low –Prevents precipitation of organic anions Mg 2+ helps solubilize organic anions

How cells keep [Ca] i low All eukaryotic cells have PM Ca 2+ -ATPase –Excitable cells also have Na + /Ca 2+ exchanger (NCX) ER Ca 2+ -ATPase (against a high grad) Mitochondrial high capacity (low affinity) pump –When [Ca] i very high (dangerous) levels (>10 -5 M) –Inner mitochondrial membrane –Uses the electrochemical gradient generated during electron-transfer of oxidative-phosphorylation

Calcium Concentrations [Ca 2+ ] o / [Ca 2+ ] i >10 4 –[Ca 2+ ] o ~10 -3 M –[Ca 2+ ] ER ~10 -3 M –[Ca 2+ ] i <10 -7 M at rest

Ca 2+ - a versatile signal Target TissueSignaling MoleculeMajor Responses LiverVasopressinGlycogen breakdown PancreasAChAmylase secretion Smooth muscleAChContraction Mast cellsAntigenHistamine secretion Blood plateletsThrombinaggregation

Ca 2+ - a versatile signal Synaptic vesicle release (  s) Excitation-contraction coupling (ms) Smooth muscle relaxation (ms-sec) Excitation-transcription coupling (min-h) Gene transcription (h) Fertilization (h)

How Ca 2+ is measured Ratiometric dyes Cameleons

How cells ↑ [Ca] i Voltage-gated Ca 2+ Channels –Membrane potential drives Ca 2+ down its chemical gradient –Different channels in different cells Different properties for different purposes

Ca 2+ shut-off pathways Voltage-gated Ca 2+ channels inactivate IP 3 rapidly dephosphorylated Ca 2+ rapidly pumped out

Biochemical Society Transactions (2003) Volume 31, part 5 Ca 2+ oscillations

Intracellular Ca 2+ Targets EF hand-containing proteins –Calmodulin C2 domain proteins –PKC –Synaptogamin

Intracellular Ca 2+ Targets EF hand proteins Troponin C Calmodulin Calcineurin Calpain Recoverin S100

Calmodulin (CaM) 16.8 kDa Ca 2+ -binding protein Highly conserved in all eukaryotes Up to 1% of protein Binds 4 mol Ca 2+ /mol Binding affinity ~.25-1x10 6 M –Cooperativity with Hill constant ~ 2

Cell, Vol. 108, 739–742, March 22, 2002 Classical mode of CaM regulation Relief of autoinhibition

CaMKII Autoinhibition and Frequency Decoder

New modes of CaM interaction Cell, Vol. 108, 739–742, March 22, 2002

New modes of CaM interaction New modes of CaM regulation

Intracellular Ca 2+ Targets C2 Domains Protein kinase C Synaptotagmin Phospholipase A 2

C2 domains - PKC Chem. Rev. 2001, 101,

Synaptotagmin and neurotransmitter release