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The Journal of Neuroscience, March 1, 2000, 20(5):1722-1734
Mechanisms of Calcium Decay Kinetics in Hippocampal Spines: Role
of Spine Calcium Pumps and Calcium Diffusion through the Spine Neck in
Biochemical Compartmentalization
Ania
Majewska1,
Edward
Brown2,
Jonathan
Ross1, and
Rafael
Yuste1
1 Department of Biological Sciences, Columbia
University, New York, New York 10027, and 2 Department of
Applied Physics, Cornell University, Ithaca, New York 14853
Dendritic spines receive most excitatory inputs in the CNS and
compartmentalize calcium. Although the mechanisms of calcium influx
into spines have been explored, it is unknown what determines the
calcium decay kinetics in spines. With two-photon microscopy we
investigate action potential-induced calcium dynamics in spines from
rat CA1 pyramidal neurons in slices. The
[Ca2+]i in most spines shows two decay
kinetics: an initial fast component, during which
[Ca2+]i in spines decays to dendritic
levels, followed by a slower decay phase in which the spine follows
dendritic kinetics. The correlation between
[Ca2+]i in spine and dendrite at the
breakpoint of the decay kinetics demonstrates diffusional equilibration
between spine and dendrite during the slower component. To explain the
faster initial decay, we rule out saturation or kinetic effects of
endogenous or exogenous buffers and focus instead on (1) active calcium
extrusion and (2) buffered diffusion of calcium from spine to dendrite.
The presence of an undershoot in most spines indicates that extrusion mechanisms can be intrinsic to the spine. Supporting the two
mechanisms, pharmacological blockade of smooth endoplasmic reticulum
calcium (SERCA) pumps and the length of the spine neck affect spine
decay kinetics. Using a mathematical model, we find that the
contribution of calcium pumps and diffusion varies from spine to spine.
We conclude that dendritic spines have calcium pumps and that their density and kinetics, together with the morphology of the spine neck,
determine the time during which the spine compartmentalizes calcium.
Key words:
CA1; imaging; two-photon; buffer; LTP; slices; SERCA; CPA
Copyright © 2000 Society for Neuroscience 0270-6474/00/2051722-13$05.00/0
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