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Intracellular calcium clearance in Purkinje cell somata from rat cerebellar slices

Leonardo Fierro, Reinaldo DiPolo * and Isabel Llano

1. The mechanisms governing the return of intracellular calcium (Ca²⁺_i) to baseline levels following depolarization-evoked [Ca²⁺]_i rises were investigated in Purkinje cell somata using tight-seal whole-cell recordings and fura-2 microfluorometry, for peak [Ca²⁺]_i ranging from 50 nM to 2 µM.

2. Ca²⁺_i decay was well fitted by a double exponential with time constants of 0·6 and 3 s. Both time constants were independent of peak [Ca²⁺]_i but the contribution of the faster component increased with [Ca²⁺]_i.

3. Thapsigargin (10 µM) and cyclopiazonic acid (50 µM) prolonged Ca²⁺_i decay indicating that sarco-endoplasmic reticulum Ca²⁺ (SERCA) pumps contribute to Purkinje cell Ca²⁺_i clearance.

4. A modest participation in clearance was found for the plasma membrane Ca²⁺ (PMCA) pumps using 5,6-succinimidyl carboxyeosin (40 µM).

5. The Na⁺-Ca²⁺ exchanger also contributed to the clearance process, since replacement of extracellular Na⁺ by Li⁺ slowed Ca²⁺_i decay.

6. Carbonyl cyanide m-chlorophenylhydrazone (CCCP, 2 µM) and rotenone (10 µM) increased [Ca²⁺]_i and elicited large inward currents at -60 mV. Both effects were also obtained with CCCP in the absence of external Ca²⁺, suggesting that mitochondrial Ca²⁺ uptake uncouplers release Ca²⁺ from intracellular stores and may alter the membrane permeability to Ca²⁺. These effects were irreversible and impeded tests on the role of mitochondria in Ca²⁺_i clearance.

7. The relative contribution of the clearance systems characterized in this study varied as a function of [Ca²⁺]_i. At 0·5 µM Ca²⁺_i, SERCA pumps, PMCA pumps and the Na⁺-Ca²⁺ exchanger contribute equally to removal and account for 78 % of the process. Only 45 % of the removal at 2 µM Ca²⁺_i can be explained by these systems. In this high [Ca²⁺]_i range the major contribution is that of SERCA pumps (21 %) and of the Na⁺-Ca²⁺ exchanger (18 %), whereas the contribution of PMCA pumps is only 6 %.

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