Data from: Aging reduces the number and function of L-type calcium channels in the membrane of cardiac pacemaker cells
Data files
Jun 05, 2026 version files 39.25 KB
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Fig10B_caveolin3_fold_change.csv
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Fig10D_caveolin3_signal_intensity.csv
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Fig10F_CaV1_2_cluster_density.csv
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Fig10F_CaV1_3_cluster_density.csv
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Fig10G_CaV1_2_cluster_area_nm2.csv
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Fig10G_CaV1_3_cluster_area_nm2.csv
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Fig2D_AP_firing_rate_bpm.csv
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Fig2E_CV_cycle_length_pct.csv
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Fig2F_DD_duration_ms.csv
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Fig2H_EDD_slope_mVms.csv
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Fig2I_LDD_slope_mVms.csv
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Fig3B_ICaL_current_density_pApF.csv
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Fig3B_ICaT_current_density_pApF.csv
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Fig4B_cell_width_tissue_um.csv
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Fig4C_cell_length_tissue_um.csv
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Fig4E_short_axis_isolated_um.csv
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Fig4F_long_axis_isolated_um.csv
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Fig4G_cell_area_isolated_um2.csv
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Fig4H_capacitance_pF.csv
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Fig5A_NCX1_fold_change.csv
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Fig5C_NCX1_area_pct.csv
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Fig5D_NCX1_particle_density.csv
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Fig5F_NCX1_PM_area_pct.csv
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Fig5H_SR_Ca_transient_amplitude.csv
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Fig5I_SR_Ca_transient_decay_s.csv
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Fig6A_CaV1_2_fold_change.csv
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Fig6A_CaV1_3_fold_change.csv
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Fig6D_HCN4_area_pct.csv
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Fig6E_HCN4_particle_density.csv
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Fig6J_CaV1_2_particle_density.csv
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Fig6K_CaV1_3_particle_density.csv
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Fig7C_CaV1_2_cluster_area_nm2.csv
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Fig7C_CaV1_3_cluster_area_nm2.csv
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Fig7D_CaV1_2_cluster_density.csv
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Fig7D_CaV1_3_cluster_density.csv
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Fig8B_NPo_channel_open_probability.csv
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Fig9B_AP_firing_rate_bpm.csv
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Fig9E_DD_duration_ms.csv
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Fig9F_EDD_slope_mVms.csv
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Fig9G_LDD_slope_mVms.csv
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Fig9H_CV_cycle_length_pct.csv
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Fig9K_ICa_fold_change.csv
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Fig9M_SR_Ca_amplitude_fold_change.csv
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Fig9M_SR_Ca_tau_fold_change.csv
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Fig9O_If_fold_change.csv
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README.md
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Abstract
Pacemaker rate declines progressively with age across mammalian species, and in humans, this decline can become pathological, often necessitating implantation of artificial pacemakers. To investigate the cellular and molecular mechanisms underlying age‑associated pacemaker slowing, we compared sinoatrial node pacemaker cells from young and old mice. We quantified action potential parameters, L‑type calcium current density, L‑type calcium channel surface expression and clustering, and single‑channel activity. In parallel, we assessed sarcoplasmic reticulum calcium load and the expression and functional activity of the sodium‑calcium exchanger. We additionally examined the effects of pharmacological activation of L‑type calcium channels and evaluated the role of caveolin as a potential regulator of age‑dependent changes in channel organization. Data were collected using a combination of electrophysiology, fluorescence microscopy, direct stochastic optical reconstruction microscopy, and western blotting. Across these approaches, we found that aging is associated with disruption of L‑type calcium channel organization and function, including reduced channel density at the plasma membrane, decreased channel clustering, and reduced channel open probability. Pharmacological enhancement of L‑type calcium channel open probability restored pacemaker rate in aged cells to levels comparable to those observed in young cells. We further identified loss of caveolar structures as a contributing mechanism to age‑associated alterations in L‑type calcium channels. Together, these data indicate that aging impairs the organization and function of L‑type calcium channels in pacemaker cells and that this dysfunction contributes to the age‑related decline in pacemaker activity.
Dataset supporting the manuscript:
Vivas O, Baudot M, Madden R, Pinon-Teal WL, Hunt MS, Choi S, Pournejati R, Flores VA, Santana LF, Moreno CM. Aging Reduces the Number and Function of L-type Calcium Channels in the Membrane of Cardiac Pacemaker Cells. Circulation Research. Submission ID: CIRCRES/2025/327894DR1.
Corresponding author: Claudia M. Moreno (morenocm@uw.edu)
Related publication DOI: [to be added upon publication]
License
This dataset is released under the Creative Commons Zero (CC0) waiver.
Summary
This dataset contains the raw numerical data underlying the figures of the manuscript. The study quantifies age-associated changes in the electrical activity, calcium handling, protein expression, and nanoscale organization of sinoatrial node (SAN) pacemaker cells. Experiments were performed on pacemaker cells isolated from young (4–6 months) and old (24–30 months) male C57BL/6 mice. Measurements span whole-cell electrophysiology (action potentials, calcium currents, single-channel recordings), calcium imaging, confocal and super-resolution (STORM) microscopy of ion channel distributions, and Western blot quantification of pacemaker proteins.
How to open these files
All data files are plain UTF-8 encoded comma-separated values (CSV). They can be opened with:
- Any text editor (Notepad, TextEdit, VS Code, etc.)
- Microsoft Excel, Numbers, LibreOffice Calc (open directly)
- R:
read.csv("filename.csv") - Python (pandas):
pandas.read_csv("filename.csv") - MATLAB:
readtable("filename.csv")
Each CSV contains two columns. Headers are in row 1; data begins in row 2.
Empty cells in any column indicate no observation at that row position (the two columns are padded to equal length for display purposes only). They do not represent missing data.
Software used for analysis
Statistical analyses were performed in:
- GraphPad Prism 10 — used for unpaired t-tests, nested t-tests, and two-tailed Mann-Whitney tests.
- R version 4.3 or later with the
lme4package — used for the linear mixed effects models in Figure 9 panels B, F, G, and H. Model formula:value ~ age * condition + (1 | mouse/cell_id). Post-hoc comparisons used Sidak correction for four pre-specified contrasts. - ImageJ (NIH) — used to quantify Western blot band intensities as area under the curve normalized to total protein.
No custom analysis scripts are required to reproduce the comparisons reported in the manuscript. Raw values in the CSV files can be entered directly into Prism (or an equivalent package) to reproduce the statistical tests.
Abbreviations and symbols
| Abbreviation | Definition |
|---|---|
| AP | Action potential |
| bpm | Beats per minute |
| Caᵥ1.2 / Caᵥ1.3 | L-type voltage-gated calcium channel, Caᵥ1.2 (α₁C) or Caᵥ1.3 (α₁D) |
| CV | Coefficient of variation |
| DD | Diastolic depolarization |
| EDD | Early diastolic depolarization |
| HCN4 | Hyperpolarization-activated cyclic nucleotide-gated channel 4 (pacemaker cell marker) |
| ICaL | L-type calcium current |
| ICaT | T-type calcium current |
| If | Funny current |
| LDD | Late diastolic depolarization |
| MβCD | Methyl-β-cyclodextrin (cholesterol-depleting agent) |
| MDP | Maximum diastolic potential |
| N | Number of animals |
| n | Number of cells |
| NCX1 | Sodium–calcium exchanger 1 |
| NPo | Number of channels × open probability |
| PM | Plasma membrane |
| SAN | Sinoatrial node |
| SEM | Standard error of the mean |
| SR | Sarcoplasmic reticulum |
| STORM | Stochastic Optical Reconstruction Microscopy |
| TIRF | Total Internal Reflection Fluorescence (microscopy) |
| WB | Western blot |
Experimental context (common to all files)
All experiments were performed on pacemaker cells or sinoatrial node tissue from male C57BL/6 mice. Two age groups were used throughout the study:
- Young: 4–6 months
- Old: 24–30 months
Electrophysiology recordings were performed at 32–34 °C. Full experimental details (cell isolation, patch-clamp, calcium imaging, STORM, Western blot protocols) are provided in the Methods section of the manuscript.
File descriptions
The dataset is organized by manuscript figure. Each CSV contains two columns; the comparison column labels are indicated below.
Figure 2 — Spontaneous action potential parameters in young vs old pacemaker cells
Perforated-patch current-clamp recordings of spontaneous action potentials from isolated pacemaker cells.
Sample sizes: Young n = 28 cells, N = 5 mice / Old n = 18 cells, N = 5 mice. Columns Young and Old are unpaired.
| File | Panel | Measurement | Units |
|---|---|---|---|
Fig2D_AP_firing_rate_bpm.csv |
2D | Spontaneous action potential firing rate | beats per minute (bpm) |
Fig2E_CV_cycle_length_pct.csv |
2E | Coefficient of variation (CV) of cycle length, computed per cell across 50 consecutive cycles as (SD / mean) × 100 | percent (%) |
Fig2F_DD_duration_ms.csv |
2F | Diastolic depolarization (DD) duration, from the maximum diastolic potential (MDP) to the action potential threshold | milliseconds (ms) |
Fig2H_EDD_slope_mVms.csv |
2H | Early diastolic depolarization (EDD) slope, from a linear regression fit to the early DD phase (MDP to approximately –55 mV) | mV/ms |
Fig2I_LDD_slope_mVms.csv |
2I | Late diastolic depolarization (LDD) slope, from a linear regression fit to the late DD phase (approximately –55 mV to threshold) | mV/ms |
Statistical test: nested t-test (cells nested within animals), GraphPad Prism 10.
Figure 3 — Whole-cell calcium currents
Whole-cell voltage-clamp recordings in response to 20-ms depolarizing pulses to –20 mV from a holding potential of –80 mV. The L-type component (ICaL) was isolated by subtracting the nifedipine-resistant current (10 μM nifedipine) from the total calcium current; the nifedipine-resistant component represents the T-type current (ICaT).
Sample sizes: Young n = 16 cells, N = 4 mice / Old n = 16 cells, N = 4 mice.
| File | Panel | Measurement | Units |
|---|---|---|---|
Fig3B_ICaL_current_density_pApF.csv |
3B | L-type calcium current density (peak Iᴄₐₗ / cell capacitance) | pA/pF |
Fig3B_ICaT_current_density_pApF.csv |
3B | T-type calcium current density (nifedipine-resistant component / cell capacitance) | pA/pF |
Statistical test: two separate nested t-tests (one for L-type, one for T-type); these are two independent hypotheses, not a multigroup comparison.
Figure 4 — Pacemaker cell size
Three complementary approaches test whether aging affects pacemaker cell size: (1) cell dimensions measured from HCN4-immunostained whole-mount tissue explants; (2) cell dimensions measured from single isolated pacemaker cells; and (3) whole-cell capacitance as an electrophysiological measure of cell surface area.
| File | Panel | Measurement | Units | Sample sizes |
|---|---|---|---|---|
Fig4B_cell_width_tissue_um.csv |
4B | Cell width (short axis) from HCN4-immunostained whole-mount tissue | μm | Young n = 37, N = 3 tissues / Old n = 39, N = 3 tissues |
Fig4C_cell_length_tissue_um.csv |
4C | Cell length (long axis) from HCN4-immunostained whole-mount tissue | μm | Young n = 37, N = 3 tissues / Old n = 39, N = 3 tissues |
Fig4E_short_axis_isolated_um.csv |
4E | Short axis of isolated HCN4-positive pacemaker cells | μm | Young n = 53, N = 5 mice / Old n = 54, N = 5 mice |
Fig4F_long_axis_isolated_um.csv |
4F | Long axis of isolated HCN4-positive pacemaker cells | μm | Young n = 53, N = 5 mice / Old n = 54, N = 5 mice |
Fig4G_cell_area_isolated_um2.csv |
4G | Area of isolated HCN4-positive pacemaker cells | μm² | Young n = 53, N = 5 mice / Old n = 54, N = 5 mice |
Fig4H_capacitance_pF.csv |
4H | Whole-cell membrane capacitance | picofarads (pF) | Young n = 22, N = 4 mice / Old n = 21, N = 4 mice |
Statistical test: nested t-test (cells nested within animals or tissues), GraphPad Prism 10.
Figure 5 — Calcium clearance machinery (NCX1) and SR calcium load
Three complementary approaches test whether the age-associated reduction in L-type current reflects changes in calcium clearance or SR load: (1) NCX1 total protein expression by Western blot; (2) NCX1 surface density by AiryScan confocal imaging; and (3) SR calcium content estimated from caffeine-induced calcium transients.
Panel A — NCX1 total expression (Western blot)
Each data point is one animal; values are fold change relative to the young group.
| File | Panel | Measurement | Units | Sample sizes |
|---|---|---|---|---|
Fig5A_NCX1_fold_change.csv |
5A | NCX1 protein abundance (old / young), Western blot. Values normalized to total protein | fold change (young normalized to 1) | Young N = 5 / Old N = 6 animals |
Statistical test (Panel A): two-tailed Mann-Whitney test (N < 10 per group).
Immunofluorescence quantification of NCX1 on AiryScan confocal images
| File | Panel | Measurement | Units | Sample sizes |
|---|---|---|---|---|
Fig5C_NCX1_area_pct.csv |
5C | Percentage of total cell area occupied by NCX1 signal | percent (%) | Young n = 30 cells, N = 3 / Old n = 15 cells, N = 3 |
Fig5D_NCX1_particle_density.csv |
5D | NCX1 particle density per unit cell area | particles/μm² | Young n = 30 cells, N = 3 / Old n = 15 cells, N = 3 |
Fig5F_NCX1_PM_area_pct.csv |
5F | Percentage of plasma membrane (PM) area occupied by NCX1 signal (HCN4 labeling used as a plasma-membrane mask) | percent (%) | Young n = 15 cells, N = 3 / Old n = 8 cells, N = 3 |
Caffeine-induced SR calcium transient
Cells were loaded with Cal-520 and exposed to a localized 20 mM caffeine puff in calcium-free external solution.
| File | Panel | Measurement | Units | Sample sizes |
|---|---|---|---|---|
Fig5H_SR_Ca_transient_amplitude.csv |
5H | Amplitude of the caffeine-induced SR calcium transient (F/F₀) | fold change | Young n = 37 cells, N = 3 / Old n = 48 cells, N = 4 |
Fig5I_SR_Ca_transient_decay_s.csv |
5I | Decay time constant (τ) of a single-exponential fit to the decay phase of the calcium transient | seconds (s) | Young n = 36 cells, N = 3 / Old n = 47 cells, N = 4 |
Statistical test: nested t-test (cells nested within animals), GraphPad Prism 10.
Figure 6 — Caᵥ1.2 and Caᵥ1.3 total expression and surface density
Figure 6 tests whether aging affects the total protein expression (Panel A, Western blot) and the plasma membrane surface density (Panels D–K, AiryScan footprint imaging) of CaV1.2 and CaV1.3 L-type calcium channels. CaV1.2 and CaV1.3 were measured in independent experimental preparations and are not paired comparisons on the same biological unit.
Panel A — Total protein expression (Western blot)
Each data point is one animal; values are fold change relative to the young group.
| File | Panel | Measurement | Units | Sample sizes |
|---|---|---|---|---|
Fig6A_CaV1_2_fold_change.csv |
6A | CaV1.2 protein abundance, Western blot | fold change (young normalized to 1) | Young N = 4 / Old N = 4 animals |
Fig6A_CaV1_3_fold_change.csv |
6A | CaV1.3 protein abundance, Western blot | fold change (young normalized to 1) | Young N = 5 / Old N = 5 animals |
Statistical test (Panel A): two independent two-tailed Mann-Whitney tests (one for CaV1.2, one for CaV1.3). N < 10 per group.
Panels D–K — Surface density by AiryScan confocal imaging
Channel surface density at the basal (footprint) plane of isolated pacemaker cells.
| File | Panel | Measurement | Units | Sample sizes |
|---|---|---|---|---|
Fig6D_HCN4_area_pct.csv |
6D | Percentage of cell footprint area occupied by HCN4 signal | percent (%) | Young n = 10 cells, N = 3 / Old n = 12 cells, N = 3 |
Fig6E_HCN4_particle_density.csv |
6E | HCN4 particle density at the footprint | particles/μm² | Young n = 10 cells, N = 3 / Old n = 12 cells, N = 3 |
Fig6J_CaV1_2_particle_density.csv |
6J | CaV1.2 particle density at the footprint | particles/μm² | Young n = 22 cells, N = 3 / Old n = 33 cells, N = 3 |
Fig6K_CaV1_3_particle_density.csv |
6K | CaV1.3 particle density at the footprint | particles/μm² | Young n = 16 cells, N = 3 / Old n = 23 cells, N = 3 |
Statistical test (Panels D, E, J, K): nested t-test (cells nested within animals), GraphPad Prism 10.
Figure 7 — Nanoscale organization of CaV1.2 and CaV1.3 clusters (STORM)
Super-resolution microscopy (Stochastic Optical Reconstruction Microscopy, STORM) in TIRF mode was used to quantify channel clusters at the basal region (footprint plane) of isolated pacemaker cells, with a Z-axis penetration depth of approximately 100 nm. CaV1.2 and CaV1.3 were imaged in separate cells from independent preparations — they are two independent comparisons, not multiple comparisons on the same biological unit.
| File | Panel | Measurement | Units | Sample sizes |
|---|---|---|---|---|
Fig7C_CaV1_2_cluster_area_nm2.csv |
7C | Mean CaV1.2 cluster area | nm² | Young n = 8 cells, N = 3 / Old n = 6 cells, N = 3 |
Fig7C_CaV1_3_cluster_area_nm2.csv |
7C | Mean CaV1.3 cluster area | nm² | Young n = 12 cells, N = 4 / Old n = 8 cells, N = 3 |
Fig7D_CaV1_2_cluster_density.csv |
7D | CaV1.2 cluster density | clusters/μm² | Young n = 8 cells, N = 3 / Old n = 6 cells, N = 3 |
Fig7D_CaV1_3_cluster_density.csv |
7D | CaV1.3 cluster density | clusters/μm² | Young n = 12 cells, N = 4 / Old n = 8 cells, N = 3 |
Statistical test: nested t-test (cells nested within animals), GraphPad Prism 10.
Figure 8 — Single-channel activity (cell-attached patch)
Cell-attached patch-clamp recordings. N·Po (number of active channels × open probability) was calculated from 50 voltage sweeps per patch.
Sample sizes: Young n = 9 cells, N = 3 mice / Old n = 11 cells, N = 3 mice.
| File | Panel | Measurement | Units |
|---|---|---|---|
Fig8B_NPo_channel_open_probability.csv |
8B | N·Po — number of channels × open probability | dimensionless |
Statistical test: nested t-test (cells nested within animals), GraphPad Prism 10.
Figure 9 — L-type calcium channel agonist (Bay K 8644) rescue experiment
Paired measurements before and after application of the L-type calcium channel agonist Bay K 8644 in the same cells. For panels B, F, G, and H, the paired repeated-measures structure was analyzed with a linear mixed effects model accounting for both the within-cell paired design and the nesting of cells within mice. Panels K, M, and O report pre-computed fold-change values (post/pre Bay K 8644) for each cell.
Sample sizes (panels B, F, G, H): Young n = 14 cells, N = 3 mice / Old n = 11 cells, N = 3 mice.
Panel E (DD duration): Young n = 14 / Old n = 10 (one old cell had no measurable DD and is excluded).
| File | Panel | Measurement | Units |
|---|---|---|---|
Fig9B_AP_firing_rate_bpm.csv |
9B | Action potential firing rate, control and after Bay K 8644 | bpm |
Fig9E_DD_duration_ms.csv |
9E | DD duration, control and after Bay K 8644 | ms |
Fig9F_EDD_slope_mVms.csv |
9F | EDD slope, control and after Bay K 8644 | mV/ms |
Fig9G_LDD_slope_mVms.csv |
9G | LDD slope, control and after Bay K 8644 | mV/ms |
Fig9H_CV_cycle_length_pct.csv |
9H | CV of cycle length, control and after Bay K 8644 | percent (%) |
Fig9K_ICa_fold_change.csv |
9K | Fold change of L-type calcium current after Bay K 8644 (post/pre) | fold change |
Fig9M_SR_Ca_amplitude_fold_change.csv |
9M | Fold change of SR Ca²⁺ transient amplitude | fold change |
Fig9M_SR_Ca_tau_fold_change.csv |
9M | Fold change of SR Ca²⁺ transient τ (decay time) | fold change |
Fig9O_If_fold_change.csv |
9O | Fold change of the funny current Iƒ | fold change |
Statistical test (panels B, F, G, H): linear mixed effects model fitted with the lme4 package in R (value ~ age * condition + (1|mouse/cell_id)), with Sidak correction for four pre-specified contrasts: (1) young-control vs young + Bay K 8644; (2) old-control vs old + Bay K 8644; (3) young-control vs old-control; (4) young + Bay K 8644 vs old + Bay K 8644.
Statistical test (panels K, M, O): nested t-test comparing fold changes between young and old.
Figure 10 — Caveolin-3 and calcium channel surface organization
This figure combines Western blot quantification of caveolin-3 abundance (panel B), confocal immunofluorescence of caveolin-3 at the cell edge (panel D), and STORM imaging of Caᵥ1.2 and Caᵥ1.3 cluster density and area after caveolar disruption with methyl-β-cyclodextrin (MβCD, 5 mM, 30 min; panels F and G).
| File | Panel | Measurement | Units | Sample sizes | Columns |
|---|---|---|---|---|---|
Fig10B_caveolin3_fold_change.csv |
10B | Caveolin-3 protein abundance (fold change old / young), Western blot. One value per animal, normalized to total protein | fold change | N = 6 young / N = 6 old animals | Young, Old |
Fig10D_caveolin3_signal_intensity.csv |
10D | Caveolin-3 immunofluorescence intensity at the cell edge (AiryScan) | integrated density (a.u.) | Young n = 8, N = 3 / Old n = 8, N = 3 | Young, Old |
Fig10F_CaV1_2_cluster_density.csv |
10F | CaV1.2 cluster density, control vs MβCD (young cells) | clusters/μm² | Control n = 22, N = 3 / MβCD n = 18, N = 3 | Control, MβCD |
Fig10F_CaV1_3_cluster_density.csv |
10F | CaV1.3 cluster density, control vs MβCD (young cells) | clusters/μm² | Control n = 20, N = 3 / MβCD n = 18, N = 3 | Control, MβCD |
Fig10G_CaV1_2_cluster_area_nm2.csv |
10G | Mean CaV1.2 cluster area, control vs MβCD (young cells) | nm² | Control n = 22, N = 3 / MβCD n = 18, N = 3 | Control, MβCD |
Fig10G_CaV1_3_cluster_area_nm2.csv |
10G | Mean CaV1.3 cluster area, control vs MβCD (young cells) | nm² | Control n = 20, N = 3 / MβCD n = 18, N = 3 | Control, MβCD |
Statistical tests:
- Panel 10B: two-tailed Mann-Whitney test (N < 10 per group at the animal level; Old group deviated from normality, Shapiro-Wilk p = 0.014).
- Panels 10D, 10F, 10G: nested t-test (cells nested within animals), GraphPad Prism 10.
